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The Congo Expedition of The American Museum
of Natural Mstory

INTRODUCTION

By HENRY FAIRFIELD OSBORN

BULLETIN

OF
THE AMERICAN MUSEUM OF NATURAL HISTORY
Vou. XXXIX, pp. xv-xxvii |

New York, August 1, 1919

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Albertville
Alima R.
Aluta
Amadi
Ambaca
Ambriz
Rents

Aruwimi R.
Avakubi

Babeyru
Bafuka
Bafwabaka
Bafwaboli
Bafwasende
Bagboro
Bahr el Ghebel
Bali R.
Bambili
Banalia
Banana
Bangweolo L.
Banzyville
Baraka
Baringa
Barumbu
Basali
Basankusu
Basoko

Bena-Dibele
Bengamisa
Benguela
Beni

Boga
Bokakata
Bokatola
Bokote
Bokula
Bokungu
Bolengi
Bolobo
Boma
Bombai
Bombimba
Bomili
Bomokandi R.
Bomu R. tile
Bondo (Djabir)
Brazzaville
Bukama
Bukoba
Bumba
Bumbuli
Bushimai R.
Busira R.
Buta

Bwado

Cabinda
Chambezi R.
Chinchoxo
Choga L.
Coquilhatville
Cuanza R.

Dekese

Difuma
Dilemba
Dilolo
Dilolo L.
Dima
(Djabir)
Djamba
Dongo
Doruma
Dungu
Dungu R.

Duque de Braganca

Duru
Duru R.

Eala
Ekwayolo
Elila R

Elipa
Elisabethville
Entebbe
Epulu R.
Etshutshu

Faradje

Fini R.

Fort de Possel
Fort Ferreira
Fort Portal
Fort St. Porto

Gaima Mt.
Gali
Gamangui
Gangura
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Golungo Alto
Goma
Gombari
Gondokoro

Huambo

Ikelemba R.
Ila

Imese
Ingende
Inkisi R.
Inongo
Inzia R.
Irebu
Irumu
Isangi
Isiro
Iteko
Itimbiri R.
Itoko
Ituri R.

Kabalo
Kabamba L.
Kabambare
Kabele L.
Kabinda
Kabompo R.
Kabonge
Kafakumba
Kafue R.
Kagera R.
Kalamba
Kalembe-Lembe
Kalonga
Kama
Kambove
Kampala
Kanda-Kanda
Kasai R.
Kasenga
Kasindi
Kasongo
Kasuku R.
Katako Kombe
Katola
Kayoyo
Kiambi
Kibali R.

G 24° H
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THE CONGO EXPEDITION
OF
THE AMERICAN MUSEUM OF NATURAL History

By Henry FAIRFIELD OSBORN

This expedition was planned with the cordial codperation of the
Belgian Government, and it seems appropriate to introduce the
series of American Museum publications with a résumé of previous
exploration and with a brief account of the large and important scientific
work which has been accomplished under the auspices of the Belgian
Government, as well as with a reference to the expeditions by explorers
under other flags.

RESUME OF PREVIOUS EXPLORATION AND PUBLICATION!
| EARLIER PeRtop (1816-1890)

The scientific exploration of the Congo Basin dates from Captain
J. K. Tuekey’s ill-fated expedition to the mouth of the River Congo in
1816; most of the members of his party, including the botanist, Christian
Smith, died of disease shortly after their arrival in Africa. The meagre
results of this first attempt were chiefly botanical.

During the next sixty years progress was very slow, although it
was a period of active geographical exploration, with which the following
names are identified: R. F. Burton (1862-1863); G. Schweinfurth
(1870); J. Monteiro (1873); the German Loango Expedition (P.
Gissfeldt, H. Soyaux, E. Pechuel-Loesche, Bastian, 1873-1876); Fr.
Naumann (1874); V. Lovett Cameron (1874); Pogge (1875-1884);
W. Junker (1876-1883); H. M. Stanley (1876-1888); M. Buchner
(1878-1880); A. von Mechow and E. Teusz (1880); H. von Wissmann
(1880-1884); R. Bohm and P. Reichard (1880-1884); Casati (1880-—
1889); H. Johnston (1882-1883); Emin Pasha (1883); H. Capello and
R. Ivens (1884); R. Bittner (1884-1886); and Emin Pasha and Fr.
Stuhlmann (1890-1891). Owing to the material difficulties encountered
at that time, data of scientific value could be gathered only incidentally,
so that in 1890 our knowledge of the Congo Basin, its inhabitants and
natural productions was still slight.

'This résumé was prepared at the request of President Osborn by Dr. Joseph Bequaert, January 2,

j 1919.

Abe

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Abu Mumbazi
Aka R.
Akenge

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Albert-Edward L.
Al Nile

Aruwimi R.
Avakubi
Babe}
Bafuka
Bafwabaka
Bafwaboli
Bafwasende

Bagb
Bahrel Ghebel
Bali R

Banana
Bangweolo L.
Banzyyille

iene
Bomokandi R,

jomu Re
Bondo (Djabir)
piers

Bukoba
Bumba
Bumbuli_
Bushimai R.

Difoma
Dilemba
Dilolo
Dilolo L.
Dima
(Djabir)
Diamba
Dongo
Doruma
Dungu
Dongu R

Duque de Braganca

Eala
Ekwayolo
Biila i.

Elipa
Elisabethville
Entebbe

Fort de Possel
Fort Ferreira
Fort Portal
Fort St. Porto

Gaima Mt.
Gali
Gamangui

Gangura
Garamba

0
Golungo Alto
Goma
Gombari
Gondokoro

Huambo

Tbembo
Tbhoko
Ikelemba R.
Tla

Imese
Ingende
Inkisi R.
Tnongo
Inaia R.

Tturi R.

Kabalo
Kabamba L.
Kabambare

Kafue R.

Kalembe-Lembo
Kalonga
Kama

Kasuku R.
Katako Kombe
Kntala

0
Kiambi
Kibali R.

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28°

THE BELGIAN CONGO

AREA OF EXPLORATION AND ITINERARY

OF THE
CONGO EXPEDITION

THE AMERICAN MUSEUM OF NATURALHISTORY

1909-1915

Miles Kilometers
oo 7

Itinerary

"Shaded part ibdicates
area shown oh main
map.

ANGE
3

"5 _m sto
Region explored

Fo Mawambi

Lokilo,

Katak) Kombe

odja 7

Braganca

ue\de

Malange

Kibanga
Kibombo
Kigoma
ikondja
ikwit
Kilo
Kilwa
Kinda
Kindu
Kinshasa
Kiri

Ki
Ki

Kisantu
Kisengwa
Kitobola
Kivu L.
Kole
Kongolo
Koto R.
Krebedje
Kuilu R,
Kutu
Kwamouth
Kwango R.
Kwangu R-
Kwengo R.

Kwilu-Djuma R.

Lado
Landana
Lends R.
Leopold II L.
Leopoldville
Lesse
Libanza
Libenge
Libokwa
Likati
Likati R,
Likimi
Likuala R,
Lindi R.
Lingunda
Lioko
Lisaka
Lisala
Lisha
Loange R.
Loango
Loangwa R.
Lobay R.
Lobito
Lodja
Lokandu
Lokelenge
Lokilo
Lokolama
Lokoro R.
Lomami R.
Lomela
Lomela R,

Luapula R.
Lubefu

Ludima
Luebo
Luebo R.
Luele R.
Lufira R.
Luilaka R.
Lukefu
Lukenie R.

Luki
Lukolela
Lukonzolwa
Lukuga R,
Lukula
Lukulu R.
Lulonga R.
Tulua
Lulua R,
Luluabourg
Lusambo
Lusana
Lushika R.
Luvua R,
Luvungi

Madondo
Mahagi
Makala
Makoua
Malagarazi R.
Malange
Malela
Malimba
Mandungu
Mangi
Manyanga
Maringa R.
Masisi
Matadi
Mawambi
Medje
Meridi
Micici
Mingi
Moanda
Mobeka
Moero L.
Mokata
Mokoange
Moliro
Molundu
Moma
Momboyo R.
Mompono
Monbongo
Mondjuku
Mondombe
Monga
Mongala R.
Monkero
Monveda
Muasnza
Mulungu
Mundjumbuli
Mushie
Musofi

Mutombo-Mukulu

Nala
Nepoko R.

Nouvelle Anyers

Novo Redondo
Nya Lukemba
Nyangwe
Nzoro R.

Opala
Oshwe
Oso R.

Panga
Pania-Mutombo

Poko R.
Ponthierville
Popokabaka
Pungo Andongo
Pusu

Pweto

Rafni
Redjaf
Rikwa L.
Romée

Rubi RK.
Ruisamba L.
Ruki R.
Rungu
Rutshuru
Rutshuru R-
Ruwenzori Mt.
Ruzizi R.

Sakania
Salonga R.
Samba
Sampwe
San Salvador
Sanga R.
Sankuru-Lubilash R.
Sankuru R.
Semio
Semliki R.
Shabunds R
iloango R.
Shuka 4
Sili
Simba
Soueh R.
St. Antonio
St. Paul de Loanda
Stanley Pool
Stanleyville
Surunga

Tabora
Panganvike L.
Thysville
Titule

Toa |

Tonji R.
Tshela
Tshicapa R.
Tshitombe
Tshofa
Tshopo R.
Tshuapa R.
Tumba L.

Ubangi R.
Uele R.
Uere

Uere R.
Uji

A
Ukaturaka
Ulindi R.
Upemba L.
Uvira

Vankerckhovenyille
Victoria L.
Victoria Nile

Waka
Walikale
Wamasa
Wamba R.
Wazimba
Wissmann Pool

Yahila
Yakoma
Yakuluku

Yokokara R.
Yongama
Yumbi

Zambesi R.
Zambi
Zobia

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XVl . Bulletin American Museum of Natural History [Vol. XX XIX

Mopern Prriop (1890-1914)

As far as Central Africa is concerned this period of twenty-five
years was one of great scientific activity, in which so many explorers and
scientists of various nationalities participated that it is practically
impossible to enumerate them all. Therefore the following account
deals with only the most important contributions. Although it has been
the constant policy of the Congo Free State and of the Belgian Colonial
Office to aid the scientific labors of all investigators, up to the present
by far the greater part of the work has been accomplished by the
Belgians.

BreLoian.—Bia-Franqui Expedition (J. Cornet and P. Briart, 1890-
1893); Em. and Marc. Laurent (1893, 1895, 1903-1904); A. Dewévre
(1895-1896); A. Cabra and Fr. Michel (1896-1903); Ch. Lemaire
(1899-1900); Commission on Sleeping Sickness (J. Rodhain, C. Pons, F.
Vanden Branden, and J. Bequaert, 1910-1912); L. Stappers (1911-1912);
E. Hutereau and J. Van der Gucht (1911-1912); A. Pilette (1912-
1913); and J. Bequaert (1913-1915).

Britiso.—J. E. Dutton and J. L. Todd (1903-1905); S. A. Neave
(1904-1908); P. H. G. Powell-Cotton (1905-1906); Boyd Alexander
and G. B. Gosling (1906); The Ruwenzori Expedition of the British
Museum (R. B. Woosnam and A. G. F. Wollaston, 1906); S.and58. A.
Neave (1907); E. Torday (1907-1909); C. Christy (1911-1914); and
Rogers (19138-1914).

FreNcH.—Du Bourg de Bozas and Dr. Brumpt (1902); A. Chevaher
(1902-1903 and 1912); and E. Gromier (1911).

GERMAN.—G. A. von Goetzen and W. von Pahoa (1894); R
Schlechter (1899); L. Frobenius (1904-1906); S. Ledermann (1906—
1908); A. F. Duke of Mecklenburg, H. Schubotz and J. Mildbraed
(1907-1908); H. Schubotz (1911); and T. Kassner (1908).

AustTrian.—T. Thonner (1896 and 1909); and F. Grauer (1908).

IraLtian.—Elena, Duchess of Aosta (1909).

Swerpisu.—A. v. Rosen and R. E. Fries (1911-1912); E. Arrhenius
(1913-1915).

AMERICAN.—Th. Roosevelt (1910).

Prior to the organization of the Congo Free State (1885), the Congo
Basin was practically terra incognita from a scientific point of view.
Only fragmentary data had been obtained by the earliest explorers,
such as Tuckey, Schweinfurth, Pechuel Loesche, Cameron, Pogge,
Capello, and Ivens. At about the time Stanley traced the course of the
Congo River, King Leopold II conceived his far-sighted project of

1919] Osborn, The Congo Expedition XVI

opening up to civilization the interior of the ‘Dark Continent’? and
from the outset liberally encouraged scientific investigation. The Congo
Section of the Brussels Exhibition of 1897 displayed the results of the
work accomplished during the previous twenty years, and its collections
formed the nucleus of the present Congo Museum at Tervueren, near
Brussels, which therefore became the center of research in this field.

By 1914 the principal scientific achievements had been published in
quarto form in the famous series of the “Annales du Musée du Congo,”
comprising by that time fifty-four parts, with about 4600 pages of
text and 660 plates. Geology, zoology, botany, and ethnology are all
well represented, and the excellence of most of the contributions, in
both substance and illustration, is a fitting witness to the high standard
set. In addition to these official reports, a literature of almost equal
importance, although scattered in various journals, developed through
the efforts of Belgian scientific societies and of the leaders in the coloniza-
tion movement. Foremost among the Belgian contributors were G.
Boulenger, J. Cornet, Ph. Dautzenberg, E. De Wildeman, L. Dollo,
Th. Durand, J. Fraipont, Ch. Kerremans, Aug. Lameere, Em. Laurent,
H. Schouteden, and C. Van Overbergh.

ORGANIZATION OF THE CONGO EXPEDITION OF THE
AMERICAN MUSEUM

For many years the late President Morris K. Jesup had entertained
the hope that an expedition from the American Museum might be sent
to the Congo. Early in 1907, preliminary plans had been discussed with
the Secretary General of the Congo Free State, Charles Liebrechts, the
negotiations being carried on through the Consul General of the Congo
Free State in Baltimore, James Gustavus Whiteley, and the Belgian
Consul in New York, Pierre Mali, who was a personal friend of President
Jesup. In May 1907, the Director of the Museum, Hermon Carey
Bumpus, went to Brussels to confer with the Belgian authorities. King
Leopold II generously gave his patronage to the project and graciously
presented to the American Museum an ethnological collection from the
Congo which now forms an important part of the African exhibits of the
Museum.

The plan was taken up again by Henry Fairfield Osborn on assuming
the Presidency of the American Museum, and late in the autumn of
1908, a Special Committee on the Congo Expedition was appointed,
consisting of Messrs. John B. Trevor, Chairman, Hermon C. Bumpus,

XVill Bulletin American Museum of Natural History [Vol. XXXIX

James Gustavus Whiteley, Robert W. Goelet, Herbert L. Bridgman and
Frank M. Chapman. The organization of this committee, and corres-
pondence carried on by President Osborn, M. Carton de Wiart, Director
Bumpus and others, gave definite form and impetus to the negotiations,
which finally secured the sanction of the Belgian Government to the
Museum’s exploration of the Congo, and the project became a reality.

The following official letters were exchanged between the Belgian
Government and the Director of The American Museum of Natural
History.

Légation de Belgique,
Washington, April 2, 1909.
Dear Mr. Bumpus:

In consequence of the conversation between yourself and Mr. Whiteley, when he
had the pleasure of meeting you in New York last January, Mr. Whiteley hastened
to write to His Excellency, the Minister of Colonies at Brussels, in order to secure
the codperation of the Belgian Colonial Administration with the scientific expedition
which the American Museum of Natural History proposes to send to the Belgian
Congo.

On my own part I also wrote to His Excellency, the Minister of Foreign Affairs
at Brussels, asking him to reeommend the proposition which Mr. Whiteley had made
to Mr. Renkin.

His Excellency, the Minister of Foreign Affairs, in reply to my letter, requests me,
in the name of his Colleague, the Minister of Colonies, to make to the American Mu-
seum of Natural History the following proposition:

The Colonial Administration, desiring to encourage the success of the contem-
plated scientific expedition in the Congo, offers the American Museum the sum of
6,800 franes as a contribution to the expenses of transportation of the American
mission in the territory of the Belgian Congo.

The Museum will engage, on its part, to give the Musée du Congo, in Belgium, a
participation in the scientific results of the Mission, by sending it specimens of differ-
ent species of animals which it has not at present in its collection, or which are needed
to complete its collections.

The annexed list contains the names of the animals (mammals and ibaa which
the Museum desires.

His Excellency, the Minister of Colonies, will take pleasure in recommnendige: the
American scientists to the good offices of the Colonial authorities, but it will, of course, _
be understood that the American Museum will have to pay the cost of the maintenance
and of the transport of the members of the Mission, and will also provide for all their
needs.

I shal! be very much obliged, my dear Mr. Bumpus, if you will be kind enough
to let me know whether the American Museum accepts this proposition, and in the
meantime I beg you to accept the assurance of my high regard.

(Signed) Baron Moncheur.!

1Baron Ludovic Moncheur was elected by the Trustees to an Honorary Fellowship in the Museum
on Mag 10, 1909.

1919] Osborn, The Congo Expedition XIX

New York City,
April 8, 1909.

To His Excellency Baron Moncheur,
Légation de Belgique,

Washington, D. C.

Dear Sir:

I am instructed by Henry Fairfield Osborn, President of the Board of Trustees of
The American Museum of Natural History, to acknowledge the receipt of your most
courteous favor of April second, conveying the information that through your kind
intervention, and also through the instrumentality of the Honorable James Gustavus
Whiteley, the attention of His Excellency the Minister of Colonies of the Kingdom of
Belgium has been called to our desires concerning scientific work in the Belgian
Colony of the Congo, and transmitting a most welcome series of propositions formu-
lated by His Excellency the Minister of Foreign Affairs in the name of His Colleague
the Minister of Colonies.

President Osborn wishes especially that the Colonial Administration of the King-
dom of Belgium should be informed that its desire to encourage the success of the con-
templated scientific expedition is, in itself, the most important of those factors which
will lead to this success. Moreover, he wishes me to say that the contribution of the
Colonial Administration—generous, as unexpected—indicates an attitude towards
scientific research which is most high-minded and which argues, for those having the
affairs of the Colony of the Congo in charge, an administration of caution, of liberality
and of wisdom.

The American Museum of Natural History will consider it a privilege to be
permitted to share the scientific results of this Expedition to the Congo with the
M usée du Congo, in Belgium, and to do everything in its power to develop the collec-
tions from the Congo, exhibited, and to be exhibited at Tervueren—indeed, the
Trustees of the American Museum desire that they may do much more than is sug-
gested by your formal list of desiderata.

With the utmost gratitude for your most efficient services, and confident that the
combined efforts of the Colonial Administration of the Kingdom of Belgium and The
American Museum of Natural History will result in the general promotion of science
and thus redound to the benefit of all people, I am,

Very respectfully yours,

(Signed) Hermon Carey Bumpus,
Director.

It will be observed in the foregoing correspondence that the Govern-
ment of Belgium contributed the sum of 6,800 francs ($1,329.13)
towards the first year’s expenses of field-work (estimated at $11,000)
and that the Museum engaged to enrich the collections of the Belgian
Colonial Museum at Tervueren. The instructions of the Belgian Govern-
ment were carried out in a most courteous and obliging manner by the
representatives of the Congo Colonial Administration throughout the
duration of the expedition.

XX Bulletin American Museum of Natural History [Vol. XX XIX

The expedition was at first financed to the extent of $10,000 through
the individual contributions of several Trustees and other friends of
the Museum, especially by Messrs. John B. Trevor, Charles Lanier,
Cleveland H. Dodge, J. P. Morgan, Jr., William K. Vanderbilt, A. D.
Juilliard, Robert W. Goelet, and William Rockefeller.

Mr. Herbert Lang! was chosen leader of the expedition and Mr.
James P. Chapin of Columbia College volunteered to go as Assistant.
At the end of the first year Messrs. Lang and Chapin reported the results,
which were far beyond the original expectations, and requested an ex-
tension of time.

The expedition ultimately extended over a period of six years, in
the course of which very full field reports were made by Mr. Lang from
time to time. The total expenditures of the expedition for field work
amounted to a very much larger sum than was originally contemplated,
namely, $58,000, which was raised as follows:

Contribution by the Belgian Government, (6,800 francs) $1,329.13
Subscriptions from the Trustees and other friendsof the Museum 29,000.00
Appropriations from the Jesup Endowment Fund 27,670.87

It is interesting to recall that the late President Jesup was originally
interested in the exploration of the Congo and that through his munifi-
cent bequest to the Museum he became the benefactor who made pos-
sible the continuation of this work. It is also through the Jesup Fund
that the Museum is enabled to issue the series of publications projected.

NARRATIVE OF THE EXPEDITION

The Museum party left New York on May 8, 1909, for Antwerp,
and, after receiving additional courtesies and assistance in Brussels,
sailed for Boma, arriving there June 22, when the work of the expedition
began, as told in the following narrative by Mr. Lang.

‘At President Osborn’s request the Expedition proceeded without
delay to the most promising zoological regions, 1200 to 1500 miles in-
land, a fact that contributed as much to the success as did the general
organization and excellent equipment. There, in the northeastern
Belgian Congo, it was hoped we could secure for the proposed African

iMr. Herbert Lang became connected with the American Museum staff in August 1903, and until
1906 worked upon the faunistic exhibits and habitat groups of North American birds. In 1906 he
represented the Museum on the Tjader Expedition to Africa, the expenses of which were chiefly borne
by Mr. Samuel Thorne. In 1907-1908, he worked on the material collected by the Tjader Expedition
and in preparation for the Congo Expedition. During the years 1909-1915 he was in charge of the
Congo Expedition. Upon his return he was assigned to the preparation, arrangement and description
of the Congo collections as Assistant in Mammalogy. On February 3, 1919, he was appointed Assis-
tant Curator, Department of Mammalogy.

1919] Osborn, The Congo Expedition XXi

Hall of the Museum the requisite material for habitat groups of the rare
Okapi and square-lipped Rhinoceros before their extermination made
this impossible On the journey up the Congo River from Leopoldville
(July 12, 1909) occasional collecting familiarized us with the more com-
mon faunal types. Arriving at Stanleyville August 3, necessary prepara-
tions for the porterage and future disposition of loads were made. A
month later, the Expedition, with a caravan of two hundred porters,
started upon the overland journey through the Rain Forest to Avakubi,
on the Ituri River, where, on September 30, we established our perma-
nent base. During the next three months, spent in the vicinity of
Avakubi, Ngayu, and Bafwabaka, the collections increased satisfactorily.
Our greatest efforts, however, were devoted to training a staff of fifteen
natives in various methods of collecting and adequately preserving the
material gathered, a measure of utmost importance in regions where the
destructive effects of the hot, moist climate had to be met. Such an
arrangement later allowed us to give more time to a zoological survey,
and on many side trips the preparation of collections could be accom-
plished with greater facility.

“From January to October 1910, with a base at Medje, we estab-
lished at least forty camps in the uninhabited rain forest south of the
Nepoko River. In October we could report that all necessary data and
material for an Okapi group had been obtained. Water-color sketches,
several hundred correlated photographs, accessories, including parts of
trees, lianas, bushes, samples of soil, and leaf moulds, supplemented by a
thorough study of the little-known life history, assured an ideal reproduc-
tion of such a group. The general collections also were successfully
increased, and reached a total of 1054 mammals, 1885 birds, 829 rep-
tiles and batrachians, 39 fishes, 15,000 invertebrates, and an ethnographi-
cal collection of over 700 specimens.

“The Museum authorities generously appropriated funds for a
continuation of the work in the savannah country of the Upper Uele,
where we established base camps at Niangara, Dungu, Faradje, Aba,
Vankerckhovenville, Yakuluku and Garamba, from January 1911-
July 1913. The square-lipped Rhinoceros proved fairly numerous here
and we were fortunate in obtaining for a habitat group a bull with a 42-
inch horn and a female with one 36.25 inches long, records from this
region and the largest complete specimens ever collected for exhibition.
Short hunting trips were made from Garamba into the Anglo-Egyptian
Sudan, with the Sirdar’s kind permission, and quite unexpectedly Giant
Eland, the largest known antelope, furnished equally splendid material

XX - Bulletin American Museum of Natural History [Vol. XX XIX

for a similar group. The total results by February 1913 speak for
themselves: 3227 mammals, 2244 reptiles and batrachians, 4488 birds,
1606 fishes, 40,000 invertebrates, and an ethnographical collection of
1900 items.

“From then on, the problem of transportation became our chief
concern. Over two hundred loads were stored in Medje, the same num-
ber in Niangara, besides specimens for several hundred more in Avakubi.
In Faradje alone over six hundred loads awaited removal to Stanleyville,
the nearest shipping center, by a sixty-five days’ march, more than half
of which led through dense forest; river transit was out of the question
since native dugouts could not be used for objects affected by water.
Restrictive measures connected with sleeping sickness had closed the
Nile route and the precarious condition of communication in the north-
eastern Uele made it necessary for the expedition to fashion from the
raw material everything needed for packing purposes. Trees were cut,
planks sawed, iron ore reduced, nails hammered, and ropes twisted,
since the collections could be transferred with safety only when care-
fully packed in boxes or other well-made parcels. Furthermore, the work
was necessarily slow, as the natives recruited in this region would not -
carry for more than six days, and in the forest only a couple of days,
before returning to their respective villages, so that during five years’
field work over 38,000 porters were engaged by the Congo Expedition.
Then, too, caravans exceeding one hundred porters would have met with
difficulties, especially in obtaining provisions.

“Under these circumstances Mr. Chapin chose to supervise the
transportation of all collections to Stanleyville and we both left Faradje
on February 19, 1913, he taking the direct road to Dungu, while I
passed northward to Yakuluku and Bafuka, gathering during the next
four months a valuable series of the rare Bongo as well as other material
that increased the importance of our data on distribution. In the mean-
time, two hundred loads had been removed from Niangara, where I
joined Mr. Chapin for a week, and on the first of July we parted com-
pany at Rungu for the next thirteen months, during which he directed
the transfer of nearly 1200 loads to Stanleyville, meanwhile adding to the
collections. I proceeded to Nala, Poko, and southward to Niapu, and
secured many desiderata, chief among which were series of rare forest
mammals. Certain gaps in the study of the Okapi were also filled in,
and a calf, intended for the New York Zoological Society, was captured
alive, but unfortunately succumbed later, owing to the lack of proper
food. I forwarded the new collections together with those which had

1919] Osborn, The Congo Expedition XXill

been stored at Medje for three years, and on July 25 I met Mr. Chapin
again near Avakubi. After vacating our base there we left the Ituri
district on September 9, passed down the Aruwimi to Banalia by native
canoe, proceeded thence by land to Bengamisa, and descended the Lindi
River to Stanleyville. We arrived at the last-named place on September
30, after an absence of five years in regions where steam whistles, tele-
graphs, telephones, and motors were unknown, although Stanleyville,
1200 miles inland, was connected with Europe and the Cape by wireless,
and under normal conditions steamers of 500 tons arrived every fort-
night. We had traveled about 15,000 miles on foot without accident or
sickness, although the unhealthy condition of the country caused the
government to reduce the term for resident officials from three to two
years.

“Mr. Chapin left Stanleyville for America on December 10, 1914
with the first large shipment of collections, and, passing through Liver-
pool and London, reached New York on March 31, 1915. I followed in
May with the last of the fifty-four tons of material, but at Matadi
marine transportation had been interrupted by the war, and not until
late in August were all of the collections safely on their way to America.
This delay gave an opportunity for a fruitful exploration of the Congo
estuary, especially in the neighborhood of Zambi, Malela, Banana and
St. Antonio. Leaving Banana for St. Paul de Loanda on September 14,
I sailed for New York, via Lisbon, arriving November 12, 1915, after
an absence of six years and a half.

“Tt is interesting to note that by 1909, the first year of the Congo
Expedition, the Congo Free State had become a Belgian Colony and
King Albert I, at that time heir to the throne and interested in the
aspirations and welfare of his people, made a tour of inspection through
the Congo Basin. At Stanleyville, September 1909, the Museum’s
expedition received important advice from various members of the royal
advance party, headed by the Minister of Colonies, Jules Renkin. At
Brussels, and throughout the Expeditions’ travels in Africa, invaluable
information and other assistance were given by the following Belgian
dignitaries and officials: Prince Albert de Ligne (Attaché); Felix
Fuchs, L. Henry (Governors General); M. Malfeyt, A. Lantonnois, A.
De Meulemeester, L. Moulaert (Vice-governors); Ed. Kervyn (Director
General); H. Droogmans, Mau. Van Damme (Colonial Secretaries) ;
G. Bertrand, Ch. Delhaise, Mau. Siffer, E. Verdick (Commissioners) ;
Ch. Smets (Judge); Dr. Van Campenhout, Dr. E. Etienne, and Dr. J.
Rodhain.

XXIV

Bulletin American Museum of Natural History

(Vol. XXXIX

“Henry Lane Wilson, American Ambassador to Belgium, and the
American Consuls General, William H. Handley and Harry A. McBride,
courteously represented and furthered the interests of the Expedition

at various times.”’

ITINERARY OF THE CONGO I}XPEDITION

1909 May 8
June 3
June 22
June 24-30
July 1-12
August 3-—September 4
September 12
September 24
September 30—December 7
December 10-26
December 27—January 10,
1910
January 13—October 15
October 18
October 23
October 26°
October 28
November 1—January 20,
1911
1911 January 25-30
1911-1913 February 6, 1911—February
19, 1913

1910

1911 July 12-18
August 9-12
November 2-6
December 10-22
March 10-16
April 7-24

May 4-July 24

February 19
March 2-4
March 12-24
June 14

February 19
February 24—March 1
March 5

June 14-21
June 24—July 1

1912

1913

Leave New York
Leave Antwerp
Leave Boma
Matadi
Leopoldville
Stanleyville
Bafwaboli
Bafwasende
Avakubi

Ngayu

Bafwabaka
Medje
Pawa

Isiro

Nala
Rungu

Niangara and vicinity

Dungu

Northeastern Uele—

base at Faradje

Aba and vicinity
Vankerckhovenville
Yakuluku

Aba and vicinity
Garamba
Vankerckhovenville
Garamba

Faradje
Yakuluku
Bafuka
Niangara

Faradje
Dungu
Niangara

Niangara
Rungu

Chapin

Lang

Lang and Chapin
Lang

Lang and Chapin
Lang and Chapin

(oa

Chapin

>Lang and Chapin

1919]

1914
1913

1914

1915

Osborn, The Congo Expedition

July 6-10

July 15—August 29

September 1—October 31

November 2—February 20,
1914

February 27—July 22

July 1

July 5-15

July 25

July 27

July 31—January 2, 1914
April 19

April 21

July 25-August 1
August 3

August 5-September 9
September 10
September 12
September 22-25
September 28
September 30

December 10
December 20
December 24
December 29
March 14
March 31

May 10

May 18-22

May 22-31

June 9-16

June 17

June 17—July 2

Tlye2= 12,

July 19-25

July 25—-August 1
August 6—September 14
September 15—October 1
October 10—November 2
November 12

SUMMARY OF THE COLLECTIONS

Nala
Poko
Akenge

Niapu
Medje

Rungu
Pawa
Bafwabaka
Ngayu
Avakubi
Penge
Epulu River

Babeyru
Ngayu
Avakubi
Bomili
Panga
Banalia
Bengamisa
Stanleyville

Left Stanleyville
Kinshasa
Matadi

Boma

Liverpool

New York

Left Stanleyville
Kinshasa
Leopoldville
Matadi

Boma

Zambi

Malela

Banana

St. Antonio
Banana

St. Paul de Loanda

Lisbon
New York

XXV

‘Chapin

Lang and Chapin

Chapin

Lang

The collections are chiefly zoological, representing nearly all
branches of natural history of the region traversed.

The following is

XXVl Bulletin American Museum of Natural History [Vol. XX XTX

an estimate of the number of specimens which by the middle of
November 1915 had reached the Museum safely, in spite of the
unsettled conditions and difficulties of transportation due to the war.

Mammalogy 5800
Ornithology 6200
Herpetology 4800
Ichthyology 6000
Invertebrates over 100,000
Paleontology was represented by only a few specimens referable to

ichthyology.

Anthropology—3800 specimens were added to the collection al-
ready presented to the Museum by King Leopold II.

The illustrative material includes about 300 drawings, in water-
color and ink, by Mr. Chapin, and a superb collection of 9890 photo-
graphs, the property of Mr. Lang, who has, however, placed them on
permanent deposit in the Museum. The latter relate to the following
subjects:

Anthropology 5461
Mammalogy 2155
Ornithology 512
Herpetology 365
Ichthyology 182
Invertebrate Zoology 294
Botany 483
Miscellaneous 438

The value of this collection is greatly enhanced by the detailed
diaries, note-books, observations and measurements taken in the field.
The records afford invaluable data for all the zoological and ethno-
logical studies to be published.

PROJECT OF PUBLICATION

The publications at present contemplated are planned in four series,

as follows:

1. Scientific Papers published in Bulletin form of the American
Museum, of which at present twelve volumes are projected,
under the title ZooLoGy oF THE BELGIAN Conco. These
papers will first be published in the Bulletin. They will
then be issued in a special edition of 150 copies as separate
volumes.

1919] Osborn, The Congo Expedition XXVI1

2. Memoirs of The American Museum of Natural History, of
which the volume on the Okapi is in course of preparation.
3. Ethnological Albums, in which the principal anthropological
results are to be brought together in three volumes.
4. Narrative of the Congo Expedition, in two volumes, by Herbert
Lang and James Chapin.
The division of the scientific material already assigned is as follows.
Mammalogy.—J. A. Allen, N. Hollister, H. Lang, J. P. Chapin, Childs
Frick.
Anatomical Studies.—H. von W. Schulte, C. Sharp, J. Kingsley.
Ornithology. —J. P. Chapin.
Herpetology.— K. P. Schmidt, G. K. Noble.
Ichthyology.—J. T. Nichols, L. Griscom, C. R. Eastman, L. Hussakof.
Invertebrate Zoology.
Vermes.—G. A. MacCallum.
Mollusca.—H. A. Pilsbry, J. Bequaert.
Crustacea.—M. J. Rathbun, H. A. Pilsbry, C. B. Wilson, W.G.

Van Name.
Myriapoda.—R. V. Chamberlin.
Insects.

Lepidoptera.—W. J. Holland.

Coleoptera.—C. W. Leng, A. J. Mutchler.

Orthoptera.—J. A. G. Rehn.

Neuropteroids.—N. Banks, J. G. Needham.

Diptera.—J. Bequaert, C. P. Alexander, J. 8. Hine. -

Hymenoptera.—W. M. Wheeler, J. Bequaert, I. W. Bailey,
Je ©. Bradley, (ks Ei: Lutz:

PROJECT OF INTERNATIONAL RESEARCH

In view of the international character of this exploration and of the
generous cooperation of the Belgian Government, it is proposed to make
the scientific results as well as the collections as effective as possible in
the dissemination of knowledge regarding the natural history and re-
sources of the Congo. Much of the zoological and botanical work will be
of real value in relation to the future economic development of this
great area of Africa.

The American Museum will begin by selecting, according to agree-
ment, a duplicate collection for the Congo Museum at Tervueren,
near Brussels, Belgium. This collection will include not only certain
of the more important mammals, birds, and reptiles which are still needed

XXVill Bulletin American Museum of Natural History [Vol. XX XIX

at Tervueren but also, so far as possible, paratypes of the new species
described in the series of volumes which will be collectively known as
THE ZOOLOGY OF THE BELGIAN Conco. Thus the Congo Museum at
Tervueren will be reinforced in the great work it has accomplished since
1897, the time of its establishment as a center of research in the zoology
and ethnology of the Congo. The same principle will apply to the dup-
lication of the American Museum photographs, observations, and
records of various kinds which may not be published.

In other words, the American Museum will endeavor, so far as
practicable, in every branch of science in which this expedition has
engaged, to extend its duplicates and documents for the benefit of its
sister institution in Belgium and for the dissemination of knowledge
through the opportunities which the Congo Museum at Tervueren
offers to the European students and investigators.

American Museum
of Natural History.
June 18, 1919.

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PUBLICATIONS

THE AMERICAN MUSEUM OF NATURAL HISTORY >

Mermorrs

Volume I. Zoology and Palxontology.

- Volumes JI-VIII. Anthropology. °
Volume IX. Zoology and Palzontology.
Volumes X-XIV. Anthropology.

Volumes II, IV, V, VII, VIII, X-XIV, and an Ethnographical Album form
Volumes I-X of the Memoirs of the Jesup North Pacific Expedition.
Mermorrs—NeEw SERIES
Volumes I and II. Zoology and Paleontology.
Volume ITI, part 1. Entomology.
BULLETIN

Volumes I-X XIV, XXV, parts 1 and 2, and XXVI-XXXIX.

ANTHROPOLOGICAL PAPERS

Volumes I-XI, XII, parts 1-5; XIII; XIV, parts 1 and 2; XV, part 1; XVI,
parts 1-3; XVII, parts 1-4; XVIII, parts 1-5; XIX, parts 1-3; XX, part 1;
XXI, part 1; XXII, parts 1-3; XXIII, parts 1-2; XXIV, parts 1-2; XXV, “ay
Es VA, one ule

MoNnOGRAPHS

A Review of the Primates. By D. G. Elliot. 3 volumes.

Hitherto Unpublished Plates of Tertiary Mammals and Permian Vertebrates. |

By E. D. Cope and W. D. Matthew.

Tue AMERICAN Musrum JoURNAL

Volumes I-XVIII. The Journal is a popular record of the progress of The
American Museum of Natural History, issued monthly from October to May.

HanpsBooxs. Numbers 1-7.
@
Gui1pE Lreartets. Numbers 1-48.

ANNUAL Reports. First (1869) to Fiftieth (1918).

A more detailed list, with prices, of these publications may be had upon applica-
tion to the Librarian of the Museum.

“Shrews Collected by the » Congo Expedition of the American
_ Museum.

B y N. Hotutster.

BULLETIN OF THE
AMERICAN MUSEUM: OF NATURAL HISTORY,
Vou. XXXV, Art. XXXV, pp. 663-680.

_ Scipntiric REsuLts of THE Conco Exprpition. Mammatoey, No. 1.

New York; October 21, 1916.

(Continued frum 3d page of cover.) :
Vou. XIII. ANTHROPOLOGY (not yet completed). rea
*Jesup North Pacific Expedition, Vol. TX. : ee ie
Part I.— The Yukaghir and the Yukaghirized Tungus. ey Waldemar Jochelson.. Pp.
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Vou. XIV. ANTHROPOLOGY. D
*Jesup North Pacific Expedition, Vol. X. ‘ :
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Part Il.— Haida Texts. By John R. Swanton. Pp. 271-802. 1908. Price, $5.40.

MEMOIRS.
New SeErR1Es, Vou. I.

Part I.— Crania of Tyranosaurus and Allosaurus. By Henry Fairfield Osborn, pp. 1-30,
pil. i-iv and text figures 1-27. 1912.

Parr II.— Integument of the Iguanodent Dinosaur Trachodon. By Henry Fairfield Osborn._
Pp. 31-54, pll. v—x, and text figures 1-13. 1912. Parts I and II are issued under one f
cover. , Price, $2.00.

Part Wie Craniometry of the Equide. By Bey Fairfield Osborn. Pp. 55-100, text
figures 1-17. 1912. Price, 75 cents.

Part IV.— Orthogenetic and Other Variations in Muskoxen, with a Systematic Review of
the Muskox Group, Recent and Extinct. By J. A. Allen. Pp. 103-226, pll. xi—xviii,
text figures 1-45, 1913. Price, $2.50.

Part V.— The California Gray Whale (Rhachianectes jake: Cope). By Roy C. Andrews.

Pp. 229-287, pll. xix—xxvii, text figures 1-22. 1914. Price, $2.00.

Part VI.— The Sei Whale (Balenoptera borealis Lesson). By Roy C. Andrews and H. von.

W. Schulte. Pp. 289-502, pill. xxvii-lvii, 1916. (In press.) : és Z

ETHNOGRAPHICAL- ALBUM.
Jesup North Pacific Expedition.

Ethnographical Album of the North Pacific Coasts of America and Asia. Part 1, pp. 1-5,
pll. 1-28. August, 1900. Sold by subscription, price, $6.00.

BULLETIN,

The matter in the ‘Bulletin’ consists of about 24 to 36 articles per volume, which
relate about equally to Geology, Paleontology, Mammalogy, Ornithology, Entomology,
and (in former yolumes) Anthropology, except Vol. XI, which is restricted to a ‘Cata- _ Ais
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Volume I, 1881-86........ Out of print Volume XVIII, Part I, 1902 Price, $2.00
i TE, £88790). ss, Price, $4.75 ni % : II, 1904 1.50
% TAT SOO OT ieee ry 4.00 2 z SLED E905). 2 .50

La es a ee a She ade ine 4.00 z " fe KALE oa etree 2.00
m4 NLSOS iy Me ste sin “4.00 “ SEX 1903s hic ie « 6.00
i VEE SOA aise edayeae Z 4.00 - 5 ROO A oe aie steer i 5.00
cs Mahe 1805 ies <7 4.00 7 SEAT TOOS a ieee eats id 5.00 -
f Wi EP ESO G SRR ice res ¥ 4.00 SN POORSNEE AS TOOG es cota esas % 6.00
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ANTHROPOLOGICAL PAPERS.
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AMERICAN MUSEUM. JOURNAL

The ‘Journal’ is a popular record of the progress of the American Museum of Natural
History, issued monthly, from October to May inclusive. Price, $1.50 a year. Yolumes
I-X VI, 1900-1916. 7

*The Anatomy of the Common Squid. By Leonard pees ees Williams. Pp. 1-87,
pll. i-ili, and 16 text figures. 1909.

*Chinese Pottery of the Han Dynasty. By Berthold Laufer. Pp. 1-339, pl. i-lxxv,
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For sale at the Museum.
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Agent, G. E. Stechert, 129 West 20th Street, New York City.

59.9,33 (67.5)
Article XXXV.— SHREWS COLLECTED BY THE CONGO EXPE-
DITION OF THE AMERICAN MUSEUM!

By N. HoOttuistTeEr.
Puates VII-XI.

The shrews collected by Herbert Lang and James P. Chapin on the
American Museum Congo Expedition number 177 specimens, of 15 species
and 3 genera. Almost one half of the species are new. This is not alto-
gether surprising when it is considered how few shrews have been described
from the Congo as compared with other parts of Africa. It nevertheless
seems remarkable that five of these new species should be members of the
small group of “naked-tailed”’ Crocidura of which only about ten: forms were
heretofore known. Five forms of Crocidura which have been recorded
from the general region are not represented in this collection. These are
Crocidura turba turba Dollman, C. t. tarella Dollman, C. poensis attila Doll-
man, C. boydi Dollman, and C. nigrofusca Matschie. Races of C. hilde-
gardee and C. fumosa, as well as representatives of several west coast
species also might reasonably be expected.

_ Owing to their damaged condition, immaturity, or other circumstances,
five specimens in the collection are not determinable and are not listed in
the present paper.

Of the localities listed below, Garamba, Faradje, Niangara, and Nala
are in the Uelle drainage. Medje, Gamangui, Bafwabaka, Babeyru,
Negayu, and Avakubi are in the Ituri valley. Lubila is situated on a branch
of the Tshopo, in the Stanley Falls district.

1. Crocidura nyanse kivu Osgood.

Plate X, Fig. 1.

1910. Crocidura flavescens kivu Oscoop, Ann. and Mag. Nat. Hist., Ser. 8, Vol. 5,
p. 370. April. (Lake Kivu, Congo.)

Three specimens, including one in alcohol, from Avakubi; one specimen
from Gamangui; and sixteen, including one in alcohol and one odd skull,

from Medje.
Most of the skins in this series are in a pale, worn, and faded condition,

' Scientific Results of the Congo Expedition, Mammalogy, No. 1.
663

664 Bulletin American Museum of Natural History. [Vol. XXXV,

but some are in fresh pelage and show the dark body coloration and dark
brownish feet characteristic of the race. The palest, most faded, specimens
have the feet light colored, like the coat of the body. ‘There is the variation
in color usual in series of Crocidura nyanse nyanse and C. n. kijabe; some
of the skins have dark colored bellies while others have not. This seems
to be individual variation and not a seasonal condition of pelage; the
variation is found in young as well as in adult examples.
The collectors record three pairs of inguinal mamme.

2. Crocidura surure eller.

1910. Crocidura surure HELLER, Smithsonian Misc. Coll., Vol. 56, No. 15, p. 2.
December 23. (Rhino Camp, Lado.)

One skin and skull from Faradje and an alcoholic specimen from Ga-
_ramba agree in all essential details with the type series of this species from
Rhino Camp, Lado Enclave.

3. Crocidura lutrella Heller.

1910. Crocidura lutrella HELLER, Smithsonian Misc. Coll., Vol. 56, No. 15, p. 4.
December 23. (Rhino Camp, Lado.)

A single skin (without skull) from Faradje is evidently of this species.
Compared with the type series from Lado Enclave, this specimen, which is
in older pelage, has a more grayish, less buffy belly.

4. Crocidura turba nilotica eller.

1910. Crocidura nilotica HELLER, Smithsonian Misc. Coll., Vol. 56, No. 15, p. 3.
December 23. (Rhino Camp, Lado.)

Three specimens, including two in alcohol, from Faradje are typical of
this form. A single alcoholic specimen from Nala, from which the skull has
been removed, is evidently of the same subspecies. It is browner than
usual for nlotica, but is the only July specimen seen.

5. Crocidura caliginea sp. nov.
(Plate VII, Bie. 1; VIM; Brest alge)
Type, No. 48555, Amer. Mus. Nat. Hist., skin and skull of adult Q (teeth moder-

ately worn and basal suture closed) collected at Medje, Belgian Congo, July 8, 1914,
by Herbert Lang and James P. Chapin. Orig. No. 2451.

1916.] Hollister, Shrews Collected by the Congo Expedition. 665

A medium sized, very dark brown, dark bellied, small footed species, with thinly
haired tail and with only a few long bristle hairs near base of tail.

‘Color — Entire head and body, above and below, dark brown, near to clove
brown; the upperparts very slightly brighter or more brownish, less smoky, than the
belly; but there is no distinct line of demarcation nor noticeable difference in the
shade of color above and below. The slightly brighter appearance of the back is
due to a faint speckling in the hair of cinnamon or deep buff. Fur short and close,
the undercolor deep neutral gray. Hands and feet brown, the digits yellowish.
Tail blackish brown, slightly paler at base below, appearing naked but sparingly
clothed with very short hairs; a very few longer hairs near base.

Skull and teeth Skull strongly built, with heavy maxillary processes. Brain-
ease short, wider than long, with sharp, angular corners and straight sides. Teeth
large; unicuspids from crown view very broad; third unicuspid distinctly larger
than second, especially broader across crown; the teeth all crowded in the row and
unicuspids overlapping. Upper last premolar (pm*) long as wide. The teeth thus
differ conspicuously from the type usual to the naked-tailed species of the dolichura
group and most resemble those of the forms in the turba group.

Measurements of type.— Total length, 125; tail vertebrae, 54; hind foot, 12 (dry,
without claws, 11.6); ear, 9. Skull: Condylo-incisive length, 21.3; condylo-basal ©
length, 20.6; greatest breadth, 9.5; maxillary breadth, 6.9; least interorbital breadth,
4.4; mandible, 11.8. Teeth: Entire upper row, 10; front of pm‘ to back of m?, 5.4;
entire lower row, 9.3.

The combination of dark color above and below, absence of long hairs
on tail except at base, small hind foot, short braincase, and large, roundish
unicuspid teeth, makes this species easy to recognize among the other
shrews known from the region. The species evidently belongs in that group
which includes poensis and batesi, and is not related closely to the other
bare-tailed species from Medje. It is represented only by the type speci-
men.

6. Crocidura jacksoni denti Dollman.

Plate, Eis. .2.

1915. [Crocidura] jlacksoni] denti Dottman, Ann. and Mag. Nat. Hist., Ser. 8,
Vol. 15, p. 516. May. (Between Mawambi and Avakubi, Congo.)

Seventy-one specimens, including nine in alcohol, from localities as fol-
lows: Avakubi, 6; Babeyru, 1 in alcohol; Bafwabaka, 1; Faradje, 5;
Gamangui, 2; Medje, 51; Nala, 3 in alcohol; N langara, 2.

There are specimens in this fine series collected in nearly all the months
of the year. As a consequence there is great diversity in color, the skins
in the dark, fresh coat and those in the reddish stages of extreme wear
contrasting greatly. Several very young examples show all the range of
variation exhibited by the adults; the first juvenile coat evidently fades

666 Bulletin American Museum of Natural History. [Vol. XXXV,

very rapidly. Owing to the great diversity of color shown in this series,
I have given these specimens special study in an endeavor to divide them
into two or more species, but have failed in this effort and am forced to
consider them all of one form. The skulls of the oldest males are naturally
somewhat larger than those of younger males and females, but after all the
range of variation is no greater than in several species of shrews from |
British East Africa, represented in the National Museum by equally large
series. |

There is also some variation in the relative size of the second and third
unicuspid teeth. This character certainly is not always a reliable one in
species of Crocidura which have these teeth normally somewhat of the same
size.

Specimens from the Uelle drainage appear paler than the average skins:
from the Ituri, but the pelages are not strictly comparable and the difference
is slight.

The collectors note three pairs of inguinal mamme. One female col-
lected at Bafwabaka, December 29, contained two large embryos.

7. Crocidura bicolor Bocage.

1890. Crocidura bicolor Bocaas, Jorn. Sci. Acad. Lisboa, Ser. 2; Voli Eepsa2e:
(Gambos, Angola.)

A single very immature alcoholic specimen, with skull removed, from
Avakubi. It apparently represents a form very much like C. 6b. woosnama
Dollman from Lake Ngami; but it would be useless to attempt an exact
subspecific determination. There is a faint stripe of brownish along the
upper side of the otherwise whitish tail; the white bristle hairs are rather
inconspicuous; hands and feet whitish. |

8. Crocidura oritis sp. nov.
(Plate Villa Big 92- Vali Bigs 2.0 20a)

Type, No. 48510, Amer. Mus. Nat. Hist., skin and skull of adult & (basal suture
closed; teeth moderately worn) collected at Avakubi, Ituri River, Belgian Congo,
July 6, 1914, by Herbert Lang and James P. Chapin. Orig. No. 2530.

Related to Crocidura maurisca Thomas, but averaging less blackish brown in
color, the underparts less richly colored, and the feet less blackish. Size about as in
maurisca, but hind foot larger and skull more robust, with heavier teeth. ‘Tail with
long bristles only at base as in maurisca and allies.

Color.— Type: Upperparts grayish brown, or dark hair brown, finely speckled

1916.] - Hollister, Shrews Collected by the Congo Expedition. 667

with lighter grayish; crown and face slightly darker, more blackish brown; underfur
rather light slate gray. Hands and feet brownish buff, the feet darker along outer
half from heel to toe. Tail uniformly dull blackish brown except near base below
where there are a few lighter buffy hairs. Underparts mouse gray, uniformly colored
from chin to tail except for an irregular wash of cinnamon buff. Side glands incon-
spicuous, about the color of the surrounding hair. No definite line of demarcation
between color of upperparts and of belly, the two shades blending over the sides of
the body.

Other specimens differ much in color from the type. A January skin from Medje
matches the type most closely; an April skin is in very bleached pelage, with irregu-
lar patches of rusty red on the upperparts. Two other skins from Medje, June
and September, are quite dark, more as in a very uniformly colored series of maurisca —
from the Victoria Nyanza region. These last two are, however, rather immature
animals, with the basal suture still open.

Skull and teeth.— Skull larger than in maurisca, about the size of that of the closely
related C. littoralis Heller from Butiaba, but slightly more robust in build and with
wider maxillary processes, rostrum, and palate. Teeth essentially as in littoralis and
maurisca, the unicuspids oval from crown view, with wide cingula and small cusps;
last upper premolar, in unworn condition, longer than wide.

Measurements of type — Total length, 159; tail vertebrae, 65; hind foot, 18 (dry,
without claws, 16.6). Skull: Greatest length, 23.4; condylo-incisive length, 23.3,
condylo-basal length, 22.6; greatest breadth, 10.2; maxillary breadth, 7.1; mandible,
12.1; upper tooth row, entire, 10.4; front of pm‘ to back of m?, 5.3; lower tooth row,
entire, 9.4.

This new species is based on five specimens, four from Medje and one
from Avakubi. It is closely related to maurisca and littoralis but may be
separated from either by the lighter color, greater maxillary breadth, and
more robust teeth. The three forms, maurisca, littoralis, and oritis, proba-
bly form an intergrading chain of subspecies. They are readily distin-
guished from Crocidura niobe Thomas by the larger hind foot and the
narrower unicuspid teeth.

9. Crocidura latona sp. nov.

(Plate VII, Fig. 3; VIII, Figs. 3, 3a.)

Type, No. 48610, Amer. Mus. Nat. Hist., skin and skull of adult < (basal suture
closed; teeth moderately worn) collected at Medje, Belgian Congo, March 17, 1910,
by Herbert Lang and James P. Chapin. Orig. No. 773.

Related to Crocidura niobe Thomas, but much more brownish (not gray) in color,
and with underparts scarcely lighter in color than back. Size about as in niobe.
Tail thinly haired and with long bristles only at base. Fur of back short.

Color.— Type: Upperparts rich, glossy bister, the nose darker; underfur nar-
rowly brownish gray at base. Underparts only slightly lighter brown than the back
and sides; lateral glands russet. Hands and feet buffy brown, very thinly haired

668 Bulletin American Museum of Natural History. [Vol. XXXV,

with brown. Tail above and below blackish brown, very slightly lighter at base
below.

Skull and teeth.— Skull in size about as in Crocidura niobe, or slightly smaller;
the general appearance much as in maurisca and littoralis, but decidedly smaller.
Teeth distinctly of the maurisca and littoralis type (the unicuspids rather narrow
with small cusps and large cingula) but second and third upper unicuspids wider
and more inclined to be circular, asin niobe. Last upper premolar longer than wide.

Measurements of type.— Total length, 135; tail vertebre, 59; hind foot, 14 (dry,
without claws, 13.2); ear, 9. Skull: Condylo-incisive length, 19.8; condylo-basal
length, 19.0; greatest breadth, 8.9; maxillary breadth, 6.1; mandible, 10.8; upper
tooth row, entire, 8.7; front of pm*to back of m?, 4.7; lower tooth row, entire, 8.2.

This species evidently is closely related to the Ruwenzori Crocidura
niobe, but differs conspicuously from that species in its almost uniform dark
brownish coloration, above and below. Besides the type there is a single
skin, without skull, in the collection from Avakubi which is doubtless of
the same form. It is an October specimen and is slightly more reddish
brown than the type, with which it agrees in general dimensions.

10. Crocidura ludia sp. nov.

(Riate Vil; Kies 4. xX Pigsu ules)

Type, No. 48566, Amer. Mus. Nat. Hist., skin and skull of adult o (teeth slightly
worn and basal suture not closed) collected at Medje, Belgian Congo, May 16, 1914,
by Herbert Lang and James P. Chapin. Orig. No. 2366.

A small species related to Crocidura muricauda Miller and C. dolichura Peters,
but coloration brownish, not gray, above and below. ‘Tail shorter than in dolichura
and skull smaller, with smaller teeth. Tail appearing naked but clothed with very
short, inconspicuous hairs; a few longer bristles at extreme base.

Color.— Upperparts bister or mummy brown, uniformly colored from nose to
tail; underfur smoke gray. Underparts hair brown, not conspicuously separated
from color of sides and back. Feet very thinly haired, buff, with brown streak along
outer side. Tail blackish brown above, lighter on basal third below.

Skull and teeth.— Skull like that of Crocidura muricauda but slightly larger, with
ereater interorbital breadth, wider, flatter braincase, and more slender rostrum;
palate wider and shorter. Teeth slightly smaller than in mwricauda or dolichura,
but otherwise similar, with second and third unicuspids of about equal size or second
rather smaller than third; unicuspids of the general mauwrisca type (oval in form, with
small cusps and large cingula) though not so extreme as in maurisca or littoralis;
last upper premolar longer than wide.

Measurements of type — Total length, 120; tail vertebre, 60; hind foot, 14 (dry,
without claws, 12.7); ear, 9. Skull: Condylo-incisive length, 18.2; condylo-basal
length, 17.5; greatest breadth, 8.2; maxillary breadth, 5.4; least interorbital breadth,
4.0; mandible, 9.2; upper tooth row, entire, 7.8; front of pm* to back of m?, 4.0;
lower tooth row, entire, 7.4.

1916.] Hollister, Shrews Collected by the Congo Expedition. 669

Three specimens of this new species of Crocidura are in the collection —
two from Medje and one from Ngayu. The topotype skin from Medje,
a female, measures: Total length, 116; tail vertebre, 53; hind foot, 12;
ear, 9. The tail of the Ngayu skin is 57 millimeters in length. All agree
almost precisely in coloration. The species apparently is close to C.
dolichura but may be distinguished by its brown coloration and shorter
tail.

The Liberian shrew described by Miller in 1900 as Myosorex muricauda'
proves to be a species of Crocidura. It was transferred by Thomas” in
1904 from Myosorex to the genus Sylvisorex. Consequently Dollman does
not include it in his recent revision of the African shrews of the genus Cro-
cidura.® 'The species is closely related to C. dolichura and should have been
placed between that species and maurisca in Dollman’s synopsis. It is
hard to understand how Thomas could have been misled by the excellent
description and figures given by Miller.

11. Crocidura polia sp. nov.
GPlate Wil bie. 6: EX, Bigs.- 25202)

Type, No. 48559, Amer. Mus. Nat. Hist., skin and skull of adult & (basal suture
closed; teeth moderately worn) collected at Medje, Belgian Congo, July 1, 1914, by
Herbert Lang and James P. Chapin. Orig. No. 2442.

A small, grayish brown, long-tailed shrew with the general proportions of C.
dolichura Peters, but with the tail heavily covered with short hairs which increase
in numbers and length at the tip to form a distinct white brush. A few scattered
longer bristle-hairs on basal third of tail. Skull and teeth much as in dolichura,
muricauda, and ludia. Pelage moderately full, the hairs at middle of back about
3.5 millimeters in length.

Color.— Upperparts uniform grayish brown, or perhaps better described as pale
fuscous with a faint sprinkling of silvery gray; sides and underparts distinctly lighter,
more grayish, but not sharply marked from color of back; lateral glands drab.
Hands and feet thinly coated with whitish hairs. Tail well coated with short hairs
which increase in length and numbers on the terminal half and produce a distinct
white pencil at tip; blackish brown above, neh lighter below, especially at
extremity, and tipped with white.

Skull and teeth The skull is remarkably like that of Crocidura ludia, just
described above; is of about the same size and general shape, but has a slightly
flatter braincase and less developed maxillary processes. It thus clearly shows the
relationship with dolichura and muricauda. The teeth are essentially as in ludia

1 Proc. Washington Acad. Sciences, Vol. 2, p. 645, December, 1900.
2 Proc. Zool. Soc. London, 1904, p. 199.
3 Ann. and Mag. Nat. Hist., Ser. 8, Vol. 15, pp. 508-527, May, 1915, et seq.

670 Bulletin American Museum of Natural History. [Vol. XXXV,

but the unicuspids are slightly less oval, more roundish, and pm‘ is slightly less
lengthened, about as long as wide. Second and third unicuspids about of equal size
but third overlaps second and appears larger from a view of the crown surface.

Measurements of type.— Total length, 130; tail vertebra, 72; hind foot, 13;
ear, 9. Skull: Condylo-incisive length, 18.2; condylo-basal length, 17.5; greatest
breadth, 8.2; maxillary breadth, 5.2; least interorbital breadth, 3.8; mandible, 9.4;
entire upper tooth row, 7.8; entire lower tooth row, 7.3.

The type and only specimen of this new species differs from all the
African forms of Crocidura with which I am familiar by its combination of
small size; long, hairy, penicillate tail, with longer bristle-hairs confined to
near base and few in number; and teeth of the dolichura group type.

12. Crocidura congobelgica sp. nov.
(Plate Vil io. 6:5 DX) o Pigs. 3:.300)

Type, No. 48512, Amer. Mus. Nat. Hist., skin and skull of adult @ (teeth little
worn) collected at Lubila, near Bafwasende, Belgian Congo, September 20, 1909,
by Herbert Lang and James P. Chapin. Orig. No. 122.

A small, unicolored, dark brown shrew with long, black, naked tail. General
external appearance much as in Crocidura latona, but skull widely different, more as
in Crocidura lutrella, with large maxillary processes and rounded unicuspid teeth, the
second and third of about equal size. Tail nearly naked except for scattered short,
close-lying hairs and a very few longer bristles at base.

Color.— Entire head and body, above and below, rich, glossy bister; the lower
sides and belly only very slightly lighter colored than the back. The upperparts are
very finely streaked with cinnamon buff. Underfur, above and below, pale smoke
gray. Whiskers long, mixed black and white. Hands and feet very thinly clothed
‘vith brown hairs, darker along outer sides. Tail dull brownish black except near
extreme base below where it is slightly lighter. |

Skull and teeth— The skull differs from those of other members of the naked-
tailed group in its comparatively great maxillary width. From above it greatly
resembles skulls of the otherwise very different C. lutrella Heller of Lado. The first
upper unicuspid is relatively larger, oval in crown pattern, with well developed cin-
gulum; second and third unicuspids about equal in size, the third squarely posterior
to, and somewhat overlapping, the second. In unworn condition these show the
small cusps and large cingula characteristic of the group. The last upper premolar
differs from that of other members of the group in its lack of conspicuous posterior
emargination, the median space betWeen it and the first molar being very small;
the length of this tooth is about equal to its breadth, instead of much greater as usual
in the group. From the skull of C. latona (which species this one most resembles in
color and external characters) the skull of C. congobelgica may be separated readily
by its larger size, much wider palate, stronger maxillary processes, and considerably
larger teeth. |

Measurements of type, followed by dimensions of an older adult female from
Medje: Total length, 133, —; tail vertebree, 59, —; hind foot, 14, 13. Skulls (the

1916.] Hollister, Shrews Collected by the Congo Expedition. 671

- occiput of the type skull is damaged): Condylo-incisive length, —, 20.7; condylo-
basal length, —, 20.0; greatest breadth, —, 9.1; maxillary breadth, 7.0, 6.9; least
interorbital breadth, 4.3, 4.2; palatal length, 8.9, 8.8; mandible, 10.7, 11.2; entire
upper tooth row, 9.1, 9.3; front of pm‘ to back of m?, 5.0, 5.0; entire lower tooth row,
8.4, 8.8.

There are only two specimens of Crocidura congobelgica in the collection,
the type from Lubila and a specimen from Medje. Owing to the fact that
these skins so closely resemble skins of C. latona, above described, it was with
some hesitation that I decided to recognize by name two distinct species.
Skulls of the two animals are so very different that no other course seems
open. Though both obviously belong to the same group of bare-tailed
species, the skull of the old adult male /atona is much smaller than a younger
female skull of congobelgica, and the other characters, as given above, point
to certain specific difference between the two forms. The peculiar colora-
tion is found, so far as I am aware, only in this group of species, and the
long, black, almost hairless tail further instantly separates the two species
from other shrews of the same general size.

. 13. Sylvisorex gemmeus irene Thomas.

1915. Sylvisorer gemmeus irene THoMaS, Ann. and Mag. Nat. Hist., Ser. 8, Vol. 16,
p. 151. August. (Kaganbah, Uganda.)

Eighteen specimens, including four young in alcohol, from Medje, and
one specimen from Faradje.

Compared with skins of typical gemmeus from Lado, these specimens
average considerably more brownish, less gray in color; but the forms are
certainly very closely related and the skulls seem indistinguishable: The
type locality of this subspecies is in southern Uganda, but Mr. Thomas has
already recorded specimens from the Uelle and Ituri valleys! The specimen
listed above from Faradje is like those from Medje and shows no approach
in color to the Lado form.

The collectors note three pairs of mamme for this shrew. They found
two large embryos in a female collected at Medje, January 18; and two
medium sized embryos in a female from the same place on January 20.

1 Ann. and Mag. Nat. Hist., Ser. 8, Vol. 16, pp. 151 and 471, August and December, 1915.

672 Bulletin American Museum of Natural History. [Vol. XXXV,

MEASUREMENTS OF SPECIMENS. OF Sylvisorex gemmeus irene FROM MeEpJE.

| : as er almese le Re | es
= |8| | 8 5 = S = =
S|3|s essen rae A eer allere he
Museum Shomer ihe'S Bole 2 is a = 6415 2)
exer ages Dig etctesr alae pl foe et = me > g Seige of
Number i i 5 ecco ce 7S & g x og tem h
o || & ne & (eo Bo] ee lee
SA Soniits | # z a e a |B
£ ce § Ss S Sue
acl a | Oo
48584 | ot /153/85| 16 | 9| 18.1 | 17-415.3817.7/9.3179178 moderately
| worn
A8590,) (| 136)\73)|" 150-9) 17.8") 17.3 115.5) 7-82) Obi 7 Saleen
48591 | o? | 155 | 83 | 15-| 9) 17.8 | 17.1) 5.4 1°78) 920 | 756 eee
48598 | o' | 147/81] 15 | 8) 17.5 | 16.8 | 5.3 | 7.6 | 8.9 | 7.4 | 6.8 | little worn
48599 | o& |148/73; 14/10) — — | 5.5 | — | 9.2 | 7.4 '| 6.7 | moderately
25600 3) 1146) 821 15.) 9) 9) 58 | 8.0 Orb eZ On mame
48585 | 9 | 1387/73/15 | 9/ 17.3 | 16.6/ 5.4] 7.4] 8.8/7.4]6.8 ;
48586 | 9 |142/80/ 14; 9} — | — |5.6| — | — | 7.8169 ‘
48589 | 2 438/74) 15.) 8| 17.3 | 16.5 | 5.4) 7.5) 88 72¢ ao i
48595 | 9 |141|76/ 14] 8/ 17.8] 17.38] 5.4] 7.5 | 9.0] 7.7 16.9 ¢
A48596.\99 4 140) 75) 15)| 9) 17.3 | 16.7 | 5:44) %e74|\-839)| ee eee é
48507 | @ 9) 151/83] 14 | 8) 17.5) 16.8) 5.7 | 7.8 | 8.6 °7 so Geom

14. Sylvisorex oriundus sp. nov.
(Plate VII, Big. 7; IX, Figs. 4; 40>" Xi) Grey i)

Type, No. 48554, Amer. Mus. Nat. Hist., skin and skull of adult @ (teeth little
worn) collected at Medje, Nava River, Belgian Congo, May 20, 1914, by Herbert
Lang and James P. Chapin. Orig. No. 2368.

A large, short tailed, short furred, dark bellied species related to Sylvisorex morio
(Gray), but larger, with much larger hind foot.

Color— Upperparts from nose to tip of tail dark olive brown, or, in certain lights,
dark sepia; the underfur deep neutral gray. Lower sides and underparts slightly
paler, dark grayish brown, very slightly lighter on throat. Tail slightly darker than
body above, lighter brown on basal two-thirds below. Hands and feet light buffy
brown, the ankles blackish.

Skull and teeth.— Skull larger than in S. morio, with longer tooth row. Compared
with the figure of the dentition of the type specimen of S. morio,! the second upper
unicuspid tooth is relatively much smaller; the front face of the large cusp of pm
is much less sloping, the heel of this tooth smaller, and the anterior cusp, though little
worn, is not more than half the height of the last unicuspid. The whole dentition is,

1 Dobson, Monog. Insect., pl. 25, fig. 2.

1916.| Hollister, Shrews Collected by the Congo Expedition. 673

except for the small size of the anterior cusp on pm’, and the presence of two conspicu-
ous notches on the anterior mandibular tooth, exceedingly like that of the much
smaller S. johnstoni (Dobson) as figured on plate 28 of the Monograph of the
Insectivora.

Measurements of type-— Total length, 134; tail vertebrze, 62; hind foot, 18;
ear, 8. The hind foot in the dry skin, without claws, measures 16. Skull and teeth:
Condylo-incisive length, 20.5; condylo-basal length, 19.9; maxillary breadth, 5.9;
least interorbital breadth, 4:2; mandible, 11.1; entire upper tooth row, 8.9; front of
pm‘ to back of m?, 4.4; entire lower tooth row, 8.4. The skull is badly damaged
in the braincase, but has been repaired so that the measurements as given are virtually
exact.

There is only a single specimen of this large Sylvisorex in the collection.
From the other species found at Medje, Sylvisorex gemmeus irene, it is readily
distinguished by its large size, short tail, and dark colored belly. Thomas
has recorded Sylvisorex morio from Medje ! on the basis of a single immature
specimen. The species represented may be the one here described.

15. Scutisorex congicus Thomas.
Plate XI, Fig. 2.

1915. Scutisorex congicus THomas, Ann. and Mag. Nat. Hist., Ser. 8, Vol. 16, p. 470,
December. (Medje, Congo.)

Thirty-seven specimens from Medje and one from Bafwabaka.

This genus heretofore was known only from two examples, the type
specimens of Scutisorex somereni and S. congicus, both described by Thomas
since 1910. ‘The Medje specimens are topotypes, the species congicus hav-
ing been named from a specimen collected at that place by Dr. Cuthbert
Christy in April, 1914.

There is considerable variation in color shown by the specimens in this
series. On some skins there is very little rusty coloration while in others
the entire body is suffused with buff and reddish brown. There is no very
great variation in size among fully adult examples, as shown by the accom-
panying table of measurements. The lambdoid crests frequently project
beyond the level of the condyles as described of S. somerent. ‘The characters
given by Thomas in separating this form from the Uganda species thus
prove to be mostly individual differences. The two forms are doubtless
very closely allied, but no specimen in the present series 1s quite so large as
the type of somerent.

This series contains specimens of all ages from very young, showing the

1 Ann. and Mag. Nat. Hist., Ser. 8, Vol. 16, p. 471, December, 1915.

674 Bulletin American Museum of Natural History. - [Vol. XXXV,

milk dentition, to old adults. The collectors record the mammez as two
pairs of inguinal; and the stomach contents of one adult male as several
caterpillars.

In making up the following table there were measured only specimens
fully adult, the skulls with the basal sutures tightly closed.

MEASUREMENTS OF ADULT SPECIMENS OF Scutisorex congicus FROM MEDJE.

| | 2 g Perse se z
« w =| ro) > | & - 2
io Ae ee Ve me LS vig eee e
[aL eb ee = B18 | 2 | Seal 2 oes
Museum | & rs i - 3s EPeO Niel A138 3| 8 OH les 2
See ® Ss SS Me ee > les) & Seige 8
Number eye Peles ileal kore Maan wlan aslo deen lies Bay pO eee
leis c= } a s | = xie | s 5 ° ®
ees a as ie ie, rb) x os & Q, = 5
ES se} = coos. eet SB | o 5 a EI 8
| = & jee avs OI Ser g =
| 2a) 7 | fa
48453 | 3 | 218 | 84 | 23 | 15 28.9 |30.3|.2 O18 916.7 | 16:7 | 83.4) G03 ies
48455 | o | 234 | 95 | 23 | 16 | 31.2 |32.4)12 § |¢.9'7.0/17.6/14.1 16.9 | 132
A8458 | 3 | 237 | 85 | 23 | 13/313 |32.1|14.0|9.2|6.9 | 1720 | 135) Gesa ees
ABAGZ | | 231. | 98 |) 23°) 15 |30.8 | 31.6] 13.2|9.1 16:7 | 17.3) Tose meme ase
48464 | of | 217 | 94 | 22 | 15 ;30.8/31.7/13,.9/9.4/6.9| 17.6) 13.8 | 7.0) 13:6
ASAGS |p) 2108 | 85)| 25, | 14 |30.1|31.6 | 13.9 9.1/6.8 | 17.3 | 1336) Geo ize
48473 | | 236 |.96 | 20°) 15 )31.1 | 32.7 | 14.319.7 | 6.9 | 1727 | 1A a ret lees
48474 | og | 238 | 92 | 24 | 15 131.3 |32.8/14.2 |9.7/6.8)17.9|14.3 | 7.1) 1a 4
48487 | # | 220 | 88 | 21 | 15 |29.9 131.1 /13.7)9.4|7.1|16.7|13.3 | 6.6) 12-6
48488 | & | 232 | 90 | 23 | 17 |30.8 [32.1 | 14.2|9.6|6.9|17.3 | 13.7 | 7.0) iam
ABA51 | 9 | 218 | 76 | 27 | 15 }29.5\ 31.2 113.2 | 9.38 614 |7 >) Tage eee lea
48471 | 9 | 206} 83 | 22 | 14 | 29.7/30:9 | 13.4 19.3 16.6 | 172 | 13.3) 7a eae
48480 | 9 | 205 | 81 | 23 | 12 |29.3|30.9113.4/9.1/6.2)16.7) 13.7 (6.8) 12N6
48489 | @ | 226 | 92 | 24 | 12 |31.0)32.3|13.7|9.2 |6.7 17-6) Toes anes sera
48486 | 2 | 243 | 95 | 23 | 16 | 31.9 |33.2 |18.7 /9.7]7.2 |18.3)14.5| 7.1) 13.5
48503 | 9 | — |— | —}) — |30.6 131.9) 14.119.5|6.5) 17.8 | TA Oi aces eae

1916.}

Fig.

cee el ees Cee

Hollister, Shrews Collected by the Congo Expedition. 675

EXPLANATION OF PLATES.

-Piate VII.

(Natural size.)

Crocidura caliginea. Type. 48555.
Crocidura oritis. Type. 48510.
Crocidura latona. Type. 48610.
Crocidura ludia. Type. 48566.
Crocidura polia. Type. 48559.
Crocidura congobelgica. Type. 48512.
Sylvisorex ortundus. Type. 48554.

Puate VIII.

(Five times natural size.)

Figs. 1, la. Crocidura caliginea. Type. 48555.
2, 2a. Crocidura oritis. Type. 48510.
3, 3a. Crocidura latona. Type. 48610.

Figs. 1

Fig. 1.

Fig 1:

Puate IX.

(Five times natural size.)

la. Crocidura ludia. Type. 48566.
a. Crocidura polia. Type. 48559.

2
3a. Crocidura congobelgica. Type. 48512.
4

a. Sylvisorex ortundus. Type. 58554.

PuaTs X.

Crocidura nyanse kivu Osgood. (About 2 natural size.) c ad., No. 48501.

Medje, June 13, 1914. Photo by Herbert Lang.
Crocidura jacksoni denti Dollman. (Slightly reduced.) @ ad., No. 48520.
Medje, May 28, 1914. Photo by Herbert Lang.

Puate XI.

Sylvisorex oriundus Hollister. Type. (Almost natural size.) @ ad., No.

48554. Medje, May 20, 1914. Photo by Herbert Lang.

Scutisorex congicus Thomas. (About } natural size.) < ad., No. 48475.

Medje, May 30, 1914. Photo by Herbert Lang.

Ae, ” a
‘SNpUNTIO XeTOSTATAG "2 ,, CLONE ee fas:
“"BOIS[OQOSU09 Oa ‘STILIO ee
“eljod BINPINOIg, °G “SIT ‘eOUISITRO BINPIOOIQ, “T “SIT
L 9 iS] v g G (

676

——
\

\ \] j

‘HOON CW ‘V Niaring

TIA StVIq “AXXX “TOA

ButitetTin A. M. N. H. Vor. XXXV, Peate VIII.

Fig. 1. Crocidura caliginea.
Fig. 2. % oritis.
’ Fig. 3. 3 latona.

677

ButtetTin A. M. N. H. Vout. XXXV, Puate IX.

Fig. 1. Crocidura ludia Fig. 3. Crocidura congobelgica.

rhe . polia. ‘4, Sylvisorex oriundus.

“3

Buwwtetin A.

Fig. 1.
Fig. 2.

Crocidura nyanse kivu Osgood.
Crocidura jacksoni denti Dollman.

679 ©
we

Vou. XXXV, Puate X.

Butitetin A. M. N. H.

Fig. 1. Sylvisorex oriundus Hollister.
Fig. 2. Scutisorex congicus Thomas.
680

melt age

eit

a
yee
Sen

>, *4
Minin
w TaN

(Continued from 4th page Pe cover. .) - ‘

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BULLETIN

THE AMERICAN MUSEUM
OF NATURAL HISTORY

VOLUME XLVII, 1922-1925

NEW YORK
PUBLISHED BY ORDER OF THE TRUSTEES
1922-1925

Pan reeres yew ty. s
Le oe B,C Se ete eas

F Se ee ne ul
; ily hee F
| . y Us & rie wie!
-- -
,
. .
\
a “ Se
ot } as

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lil

SCIENTIFIC STAFF

(As of February 28, 1925)

Freperic A. Lucas, Sc.D., Honorary Director
Grorce H. SHerwoop, A.M., Acting Director and Executive Secretary
Rosert C. Murpny, Sc.D., Assistant Director (in Scientific Correspondence,
Exhibition, and Labeling)
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I. DIVISION OF MINERALOGY, GEOLOGY, AND GEOGRAPHY
W. D. Martuew, F.R.S., Curator-in-Chief

GEOLOGY AND INVERTEBRATE PALHONTOLOGY

——__—_ ——_—__—__, Curator
CuestierR A. Reeps, Ph.D., Assoc. Curator of Invertebrate Paleontology (In Charge)
Epwarp J. Foy ss, B.S., Assistant

MINERALOGY

HERBERT P. WHITLOCK, C.E., Curator
GrorceE F. Kunz, Ph.D., Research Associate, Gems

VERTEBRATE PALAONTOLOGY

W. D. MatruEw, Ph.D., Curator-in-Chief

HENRY FarrFIELD Ossporn, LL.D., Sc.D., Honorary Curator
WALTER GRANGER, Associate Curator of Fossil Mammals
Barnum Brown, A.B., Associate Curator of Fossil Reptiles
Cuartis C. Mook, Ph.D., Associate Curator

WitiiaM K. Grecory, Ph.D., Associate in Paleontology
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LOWER INVERTEBRATES

Roy W. Miner, Ph.D., Curator

WILLARD G. Van Nang, Ph.D., Assistant Curator

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ENTOMOLOGY

Frank E. Lurz, Ph.D., Curator

A. J. Mutcutsr, Assistant Curator of Coleoptera

Frank E. Watson, B.S., Assistant in Lepidoptera

Cuar.es W. Lene, B.S., Research Associate in Coleoptera
HeEerBeErt F. Scuwarz, A.M., Research Associate in Hymenoptera
Wituiam M. WHEELER, Ph.D., Research Associate in Social Insects

ICHTHYOLOGY

BasHFoRD DgEAn, Ph.D., Honorary Curator

Joun T. Nicnots, A.B., Associate Curator of Recent Fishes
E. W. Gupecer, Ph.D., Associate in Ichthyology

Cuar_Les H. TownsEnp, Se.D., Research Associate

HERPETOLOGY
G. K. Noss, Ph.D., Curator

ORNITHOLOGY

Frank M. Cuapman, Sc.D., Curator-in-Chief

W. DeW. Miter, Associate Curator

Rosert CusHMAN Murpnuy, Sc.D., Associate Curator of Marine Birds

JaMEs P. Cuapin, Ph.D., Assistant Curator, Birds of the Eastern Hemisphere
LupLow Griscom, A.M., Assistant Curator

JonaTHAN Dwicut, M.D., Research Associate in North American Ornithology
Mrs. Eusiz M. B. Naumpere, Research Assistant

MAMMALOGY

H. E. Antuony, A.M., Associate Curator of Mammals of the Western Hemisphere
HERBERT Lang, Assistant Curator of African Mammals
Cart E. AKELEY, Associate in Mammalogy

COMPARATIVE AND Human ANATOMY

WiuiaM-K. Grecory, Ph.D., Curator

S. H. Cuuss, Assistant Curator

H. C. Raven, Assistant Curator

J. Howarp McGrecor, Ph.D., Research Associate in Human Anatomy
Duprey J. Morton, M.D., Research Associate

III. DIVISION OF ANTHROPOLOGY
CLARK WIssLER, Ph.D., Curator-in-Chief

ANTHROPOLOGY

CuLarkK WisstEr, Ph.D., Curator-in-Chief

Puiny E. Gopparp, Ph.D., Curator in Ethnology

N. C. Neuson, M. L., Associate Curator of Archeology

CuHaAr ES W. Mean, Assistant Curator of Peruvian Archeology

Louis R. Suniivan, Ph.D., Assistant Curator of Physical Anthropology

J. ALDEN Mason, Ph.D., Assistant Curator of Mexican Archeology

CuarRENCE L. Hay, A.M., Research Associate in Mexican and Central American
Archeology

Mito Hetiman, D.D.S., Research Associate in Physical Anthropology

Scientific Staff Vv

vi Scientific Staff

COMPARATIVE PHYSIOLOGY
Raupx W. Tower, Ph.D., Curator

CoMPARATIVE ANATOMY
WiiuiaM K. Grecory, Ph.D., Curator

IV. DIVISION OF ASIATIC EXPLORATION AND RESEARCH

Roy C. AnprEews, A.M., Curator-in-Chief

WALTER GRANGER, Associate Curator in Paleontology

FREDERICK K. Morris, A.M., Associate Curator in Geology and Geography
CHARLES P. Berxkry, Ph.D., [Columbia University], Research Associate in Geology
AmapbeEvus W. Grasau, 8.D., [Geological Survey of China], Research Associate
CuirForp H. Pops, Assistant in Zoology

V. DIVISION OF EDUCATION, BOOKS, PUBLICATION,
AND PRINTING

Grorce H. SHerwoop, A.M., Curator-in-Chief

LIBRARY AND PUBLICATIONS

Raupu W. Tower, Ph.D., Curator-in-Chief
Ipa Ricuarpson Hoop, A.B., Assistant Librarian

Pusiic EDUCATION

GrorGE H. SHERwoop, A.M., Curator-in-Chief
G. CiypE FisHer, Ph.D., Curator of Visual Instruction
GRACE FISHER Ramsgy, Assistant Curator

PusLic HEALTH.

CHARLES-EDWaRD Amory WINSLOW, D.P.H., Honorary Curator
Mary Greia, Assistant

ASTRONOMY
G. CLyp& Fisuer, Ph.D., in Charge

CONTENTS OF VOLUME XLVII

a EE SASL. 2 RA Sopa cr a ent Aen wean Ae
IE et IPI ARES No ee Oe ae aia ela ageldcbiayicheun, mew See =
ie ete aION OF OCPATALCS.... 30... <0... 0 ec ee cee een els
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Art. I.—The American Museum Congo Expedition Collection of Insectivora.
By A. Alien. (Plates I-IV, 1-text-figure.).. 0. . 0. eae ee tas
IIl.—Sciuride, Anomaluridez, and Idiuridze Collected by The American
Museum Congo Expedition. By J. A. Allen. (Plate V.)..........
I1J.—Carnivora Collected by The American Museum Congo Expedition.

By J. A. Allen. (Plates VI-LX XVIII, sixty-seven text figures, one

CLIT Le a OR Ba ae er ON GUE Lr TS
1V.—Primates Collected by The American Museum Congo Expedition.

By J. A. Allen. (Plates LX XIX-CLXVII, three text-figures, one

TLE [0s ince oe ale BaP nue hare oe ns a ae Age oc FA ec

283
501

DATES OF PUBLICATION OF SEPARATES

The edition of separates is 300 copies, of which about 100 are mailed on the date
of issue, and the others placed on sale in the Library.
Art. I, July 20, 1922.
«II, October 27, 1922.
“III, April 11, 1924.
“* IV, February 6, 1925.

Vill

LIST OF ILLUSTRATIONS

PLATES

Doctor Joel Asaph Allen. Frontispiece.

I.—Rhynchocyon stuhlmanni claudi, showing two color phases.

II.—Crocidura jacksoni denti; and Sylvisorex oriundus, type.

III. —Scutisorex congicus.

IV.—Crocidura nyanse kivu.

V.—Idiurus langi, new species; in color.

VI.—Thos anthus soudanicus: adult male and female.

VII.—Thos anthus soudanicus and Mellivora capensis cottont.

VIII.—Lutra maculicollis: adult female and young male.

IX.—Micraonyzx cinerea and Aonyx capensis: stuffed skins.

X.—Aonyx capensis: adult and young males.

XI.—Habitat of Lutra maculicollis and Aonyx capensis along the Bima River.

XII-XIII.—Civettictis civetia orientalis: adult male; melanistic and normal coloring
in the young; ventral view of genitalia, musk pouch, and anus.

XIV-XV.—Genetta pardina fieldiana and Genetta servalina: adult male and female;
side view of heads.

XVI.—Genetta victoriex: adult male.

XVII.—Genetta victoriz and Poiana richardsoni ochracea: side view of heads.

XVIII.—Osbornictis piscivora: type, adult male, in color.

XIX.—Habitat of Osbornictis piscivora, new species, along one of the larger forest
brooks-at the height of the rainy season.

XX.—Poiana richardsoni ochracea and Nandinia binotata.

XXI.—Nandinia binotata: adult and young males.

XXII.—Her pestes ichneumon funestus: two views of head.

XXIII.—Her pestes ichneumon funestus and Galerella ochracea ochracea,

XXIV.—AHelogale hirtula robusta, new subspecies; and Mungos gothneh.

XXV.—Crossarchus alexandri: views of adult male and head.

XXVI.—Ichneumia leucura ibeana: views of subadult male and head.

XXVII. Xenogale microdon, new species: views of adult male and head.

XXVIII.—Atilax robustus; and Atilax macrodon, new species.

XXIX.—Habitat of Atilax along shore of the Bima River.

XXX-XXXI.—Bdeogale nigripes: adult male; subadult female; side views of heads.

XXXII.—Crocuta crocuta fortis, new subspecies: adult female and young male.

XXXII-XXXIV.—Crocuta crocuta fortis, new subspecies: type skull, palatal view;
palatal view of two adult skulls, showing varying position of second upper
premolar.

XXXV.—Crocuta crocuta fortis, new subspecies, and C. c. germinans: comparative
views of right auditory bulla.

XXXVI.—Crocuta crocuta fortis, new subspecies, and C. c. germinans: comparative
views of braincase.

XXXVII-XXXVIII.—Leo leo azandicus, new subspecies: type; adult male; adult
female.

XXXIX-XL.—Leo leo azandicus, new subspecies: type skull, lateral, palatal, and
dorsal views; lateral view of mandible.

XLI.—Panthera pardus chui; and Panthera pardus iturensis, new subspecies.

x Illustrations

XLII-XLUI.—Panthera pardus suahelica: flat skins.

XLIV-LIV.—Panthera pardus chui: flat skins, showing individual variation in mark-
ings in adult males, females, and young.

LYV-LXIV.—Panthera pardus tturensis, new subspecies: flat skins, showing individual
variation in markings in adult males, females, and young.

LXV-LXVI.—Leptailurus serval faradjius, new subspecies: type, showing general
body proportions, and two views of head.

LXVII-LXVIII.—Leptailurus serval faradjius, new subspecies: flat skins, showing
individual variation in markings.

LXIX-LXX.—Leptailurus serval faradjius, new subspecies: type skull; lateral,
palatal, and dorsal views; lateral view of mandible.

LXXI.—Leptailurus serval kempi: flat skin.

LXXII-LXXIII.—Leptailurus ogilbyi pantasticta: flat skins, showing individual
variation in markings in adult females.

LXXIV.—Leptailurus serval faradjius, new subspecies; and Profelis aurata cottoni.

LXXV.—Profelis aurata cottoni: adult female; two views of head.

LX XVI.—Felis ocreata rubida: half-grown male and adult female.

LXXVII-LXXVIII.—Felis ocreata rubida: flat skins, showing individual variation in
markings.

LXXIX-LXXX.—Perodicticus potto faustus: adult male, showing habitual positions
when climbing or resting in trees, and awkward manner of progressing on the
ground; young male, clinging to a native’s hand as it clings to its mother’s
back.

LXXXI.—Galagoides demidoffit medius: adult female, showing general body propor-
tions and two views of head; young male.

LXXXII.—Papio doguera heuglini: adult male, side and front views of head.

LXXXIII.—Papio doguera tessellatus: young male, showing characteristics of
baboons—the lengthened head, peculiar form of ears, large ischial callosities,
and habitual upward bend of the tail at its base; and Lasiopyga pygerythra
griseisticta feeding on guava.

LXXXIV-LXXXV.—Cercocebus agilis: adult male; young female; three views of
head of adult male.

LXXXVI-LXXXVII.—Cercocebus albigena ituricus: adult male; young male;
three views of head of adult female.

LXXXVIII.—Cercocebus aterrimus: three views of head of subadult male.

LXXXIX.—Cercocebus aterrimus, subadult male; and Rhinostigma hamlyni, sub-
adult female.

XC.—Rhinostigma hamlyni: three views of head of subadult female.

XCI-XCII.—Lasiopyga brazze uelensis: adult male; young male; three views of
heads of adult males.

XCIII.—Lasiopyga Vhesti Vhesti: three views of head of adult male.

XCIV.—Lasiopyga Vhesti Vhesti, adult male; and Lasiopyga leucampyx stuhlmanni,
adult male.

XCV.—Lasiopyga leucampyx stuhlmanni: three views of head of adult female.

XCVI.—Lasiopyga denti: three views of head of adult male.

XCVITI.—Lasiopyga denti, adult male; and Lasiopyga ascanius cirrhorhinus.

XCVITI.—Lasiopyga ascanius cirrhorhinus: three views of head.

XCIX.—Lasiopyga cephus: adult male; two views of head.

I llustrations xl

C.—Lasiopyga pygerythra griseisticta: adult male; three views of head.

ClI.—Erythrocebus patas pyrronotus: adult male; two views of head.

CII-CIII.—Colobus langi, new species: type skull; lateral, dorsal, palatal, front, and
occipital views; crown and lateral views of mandible.

CIV.—Colobus powelli powelli: adult female; three views of head.

CV.—Colobus powelli powelli, adult female; and Colobus powelli brunneus, adult male,
showing anal region. .

CVI.—Colobus powelli brunneus: three views of heads of adult male and female.

CVII-CVIII.—Colobus abyssinicus ituricus: adult male; young male; two views of
head of adult male.

CIX. Colobus angolensis cottona and Colobus abyssinicus ituricus: three-quarter
view of heads.

CX-CXI.—Colobus angolensis cottonz: adult female; two views of head; general
body proportions of adult female; young female.

CXII-CXXI.—Colobus powelli powelli: adult male and female skulls; comparative

left lateral, dorsal, palatal, front, and occipital views; comparative left lateral
and crown views of mandible. .

CXXITI-CXX XI.—Colobus powelli brunneus: adult male and female skulls; compara-
tive left lateral, dorsal, palatal, front, and occipital views; comparative left
lateral and crown views of mandible.

CXXXII-CXLI.—Colobus abyssimicus ituricus: adult male and female skulls; com-
parative left lateral, dorsal, palatal, front, and occipital views; comparative
left lateral and crown views of mandible.

CXLI-CLI.—Colobus angolensis cottoni: adult male and female skulls; comparative
left lateral, dorsal, palatal, front, and occipital views; comparative left lateral
and crown views of mandible.

CLII-CLXV.—Pan schweinfurthii: general body proportions of adult female and
young male; front, side, and three-quarter views of heads of adult, subadult,
and young males and females.

CLXVI.—Hands and feet of Colobus angolensis cottoni and Pan schweinfurthii.

CLXVII.—Nest of Pan schweinfurthii.

Text FIGURES

PAGE
Development of dentition in Rhynchocyon stuhlmanni claudi................ 34
Lutra maculicollis: skull of adult male; lateral and palatal views; lateral view of
left mandible; crown view of left lower dentition...................... 86
Lutra maculicollis: dorsal view of skull of adult male...................... 87
Luira maculicollis: skull of juvenile male; lateral, palatal, and dorsal views;
lateral view of left mandible; crown view of left lower dentition.......... 89
Rhinarium of Lutra maculicoliis and Aonyx capensis...........0 00.0000 eee 91
Palmar surface of left fore foot and plantar surface of left hind foot of Lutra
maculicollis, Aonyx capensis, and Micraonyx cinerea.................. 93
Micraonyx cinerea: skull of adult female; lateral and palatal views; lateral
view of left mandible; crown view of left lower dentition.............. 96
Micraonyx cinerea: dorsal view of skull of adult female................... 97

Aonyx capensis: skull of adult female; lateral and palatal views............ 102

xXil Illustrations

Aonyx capensis: skull of adult female; dorsal view; lateral view of left
mandible; crown view of left lower dentition........................
Aonyx capensis: skull of juvenile male; lateral and palatal views; lateral view
of left mandible; crown view of left lowerdentition....................
Aonyx capensis: dorsal view of skull of Juvenile male....................4..
Civettictis civetta orientalis: skull of young adult male; lateral and palatal
views; lateral view of left mandible; crown view of left lower dentition. .
Civettictis civetta orientalis: skull of senile male; lateral and palatal views;
lateral view of left mandible; crown view of left lower dentition..........
Civettictis civetta orientalis: comparative dorsal views of skulls of young adult
male and senile male. iso... ore a ee
Civeitictis civetta orientalis: palmar surface of left fore foot; plantar surface of
left; hind fo0te52 0.5.0.0. BEE ee et Pa
Genetta pardina fieldiana: palmar surface of left fore foot; plantar surface of
lefithind: f00b.0) we i a oe
Diagrams of palmar surface of left fore foot and plantar surface of left hind foot
as found in genets... apne ROO
Genetta servalina: sane view oF penal if adult cael, showiners supernumerary
molar *(m?) on each side of Upper jaw.)... 5.3.22... ee
Genetia servalina: palmar surface of left fore foot; plantar surface of left hind

Rhinarium of Genetta pardina fieldiana, Genetta servalina, and Poiana richard-
SOME OCMMOCEO 6. ooo oes ob cae ee One Ped ha en oe ee
Osbornictis piscivora, new species: type skull; lateral, palatal, and dorsal views
Genetta victoriz: skull of adult male; lateral, palatal, and dorsal views........
Comparative views of Osbornictis piscivora, new species, and Genetta victoriz:
lateral view of left mandible; crown view of left lower dentition; left half
of dentition im occlusion; rhinarium...:.....:...5.2-.05 2.
Osbornictis piscivora, new species; and Genetta victoriz: palmar surface of left
fore foot; plantar surface of left hind foot...-:.. 2... 072 seen
Comparative views of right caleaneum and of dorsal surface of right astragalus
of Osbornictis piscivora, new species, and Genetta victori#.............+-.
Anterior view of distal portion of right humerus and external view of right
radius and ulna of Osbornictis piscivora, new species, and Genetia victoriz. .
External view of right scapula of Osbornictis piscivora, new species, Genetta
victoriz, and Nandinia binotata.. 00) econ. noe one ee ee eee
Osbornictis piscivora, new species, and Genetta victoriz: left lateral view of axis;
left lateral view of first three dorsal vertebre..............---.-0-eeee-
Osbornictis piscivora, new species, and Genetta victoriz: left lateral view of pelvic
region with the last lumbar, the sacral, and the first caudal vertebrz in place
Poiana richardsoni ochracea: skull of subadult male; lateral, palatal, and dorsal
views; lateral view of left mandible; crown view of left lower dentition. .
Poiana richardsoni ochracea: palmar surface of left fore foot; plantar surface
of eft hind Toot... sce Whe on a ls TH eH
Nandinia binotata: skull of adult male; lateral, palatal, and dorsal views......
Nandini binotata: adult male; lateral view of left mandible; crown view of left
lower: dentition... 2.5 tuiend ad Fah. Sods EO ee

103

104
105

110

111

Illustrations

Nandinia binotata: palmar surface of left fore foot; plantar surface of left hind

SBD a2 sw hod nese CHASE, Mathes anges Bite sh noel RR en anni) Dea SMa mn Pe eee eee ey
Calogale nyula: skull of adult male; lateral, palatal, and dorsal views; lateral
view of left mandible; crown view of left lower dentition................
Calogale nyula: palmar surface of left fore foot; plantar surface of left hind foot;
a SU MRSIDEIIOURNG eA se twa Riga Sa N oar Sod wlophe Pye fein Paatls Mya ohle Sie a, SHRINE NRL SONNE
Her pestes ichneumon parvidens: skull of adult male; lateral, palatal, and dorsal
ae ak Pe A WOT GR Maclay Ponte ice) 2, EH 244 ell 2 yoke Sich -« md Sea (aes SOAS ws F
Herpesies ichneumon parvidens: adult male; lateral view of left mandible;
Geammmview of left lower. dentition.) ois. scetuiswarceds eo.ceveeeis

Herpestes ichneumon funestus: palmar surface of left fore foot; plantar surface
RIM AM NERIELENS BEOQ eee cop 8 Sete deca) Soda ga RP cabs “assess, WA tad eS Ske MN a
Rhinarium of Herpestes ichneumon paryidens, Crossarchus alexandri, Galerella
ochracea ochracea, Helogale hirtula robusta, Mungos gothneh..............
Galerella ochracea ochracea: skull of adult male; lateral, palatal, and dorsal
views; lateral view of left mandible; crown view of left lower dentition. .. .
Galerella ochracea ochracea: palmar surface of left fore foot; plantar surface
ME MEROBLOIAG TERE yrs ie es ee LE SiS) Jes PIP ee al ve Re
Helogale hirtula robusta, new subspecies: skull of adult male; lateral, palatal,
and dorsal views; lateral view of left mandible; crown view of left lower
dentition........ RM Riera ts ent ot St Re, ME) ae. hey ahi
Helogale hirtula robusta, new subspecies: palmar surface of left fore foot; plantar
SULEELDE D2b SIRS, LONI VOTO) eR na a ee rr 2 ae
Mungos gothneh: skull of adult female; lateral, palatal, and dorsal views......
Mungos gothneh: adult female; lateral view of left mandible; crown view of
left lower eentition ME yr WAL SSI Sich adi (shard v AoaCeromtinne ors RUNS EERE

Crossarchus alexandri: skull of adult male; aoe palatal, and dorsal views. .
Crossarchus alexandri: adult male; lateral view of left mandible; crown view
DE LEP) DUST GIS S10) A era De ee arn eer nin
Crossarchus alexandri: Palaiar: si surface of left fore foot; plantar surface of left
mM I tan el eis eis a 4, wk aces eC RRS ace eee
Ichneumia leucura ibeana: skull of adult male; lateral, palatal, and dorsal

Ichneumia leucura ibeana: adult male; lateral view of left mandible; crown
Rea OMACILy ORV CL CETIUIGIOM <<. 5 <'s)<-s ays eosrci's w avecs + 3° 4.0 «os eyes Alte ee operons
Ichneumia leucura ibeana: palmar surface of left fore foot; plantar surface of
Rese INN LOMO Uy Pe TSP AB UUNIA | 28 co). cow 5 6 cl eid dlieks 0 alae « oca dlu Bo Oe oleremie ieee e
Xenogale microdon, new species: type skull; lateral, palatal, and dorsal views. .
Xenogale microdon, new species: type, adult male; lateral view of left mandible;
Gawmavmicwgor ters lower dentition. ....... 6060s boas oe cet dean e sees
Xenogale microdon, new species: palmar surface of left forefoot; plantar surface
‘Of LES na] V0 hsen see eco g ces Gas ae eee eared Onis Gay ooh nce.
Rhinarium of Xenogale microdon, new species, and Atilax macrodon, new species
Atilax macrodon, new species: type skull; lateral, palatal, and dorsal views. . .
Atilax macrodon, new species: type, adult male; lateral view of left mandible;
crown view of left lower dentition; lingual view of first lower molar.....

xiii

152

159.

162

172

173

174

178

180

181

184

185
187

188

189
190

191

192

196

197

198
200

201
203

204
206

XIV Illustrations

Atilax macrodon, new species: palmar surface of left fore foot; Pa surface a
of left hind net yebhe o ya 0d bee atd oP Me tle Wks BO Me oe ee ee 209

Bdeogale nigripes: skull of adult female; lateral, palatal, said ice views..... 210
Bdeogale nigripes: adult female; lateral view of left mandible; crown view
of left: lower dentition: .. 02... 0.02 is. CO ee 211
Bdeogale-nigripes: palmar surface of left fore foot; plantar surface of left hind
foot; rhinarium... 00.000 eee ee ee eee 212
Allenopithecus nigroviridis: skull of adult male; lateral and palatal views...... 420
Allenopithecus nigroviridis: mandible of adult male; right lateral view; pos-
terior view of right ascending ramus.....2. <<: 22). 92S. ee 421
Allenopithecus nigroviridis: dentition of adult male; crown view.............. 421
Maps
PAGE

Map of the Congo and Lake Region of Africa, showing localities where carni-
vores were collected by the Congo Expedition, as well as others mentioned
in the present. Paper's oi. 6 cs Ga de css sw ew ees ee 80
Map of the Congo and Lake Region of Africa, showing principal localities where
primates were collected by the Congo Expedition, as well as others men-
tioned inthe present. paper ::...:)..s6..:05.5..308 )40s 498

LIST OF NEW TAXONOMIC NAMES IN THIS VOLUME

GENERA

eens Sens; lo replace Le pionye .. 626 66a od ee ld ae eas i ks
Meena SM NOTIN Ante ain coed, <1. ss ar hs ir sta doe wis hago Vie Dw be

Ge BE OEE By Se. 5 SOR EVE ih aes ee RT ae eR
Conopubecus Allen. To replace Hamadryas..... 2.0.0.0... 2 cece eb eee es

SNORE OCS MR AINE Sees NE ote ahh oy ne lars orci nee hse segs ANE rake Haseinuasa teamel

RP MMIC Stee NLC 2500 ccs ys, Soho ae Sn be gh A Ae Ns he eee il
(APP Ode LIS GAIUS TR ORS a aR a a aia owe
Crocidura caliginea Hollister... .. PPR aA Cpe Aa Ones Pe eae Sg BRN os a nner Te
Crocidura oritis Hollister........ Se eres Aaa kts ae ARE x Bt eRe NS

LUNES LO, PDDUICCs 1 (CO) US i eA a
Guacnagecangoucigica. Hollister..: 2.2.2.0)... 0. ek co ee oe oe ory:
Mrmr Pens FIOMIStCR 2) 22 ee OS ea Se ed
Picvascrmmus rujoprachium medjranus Allen... 2.2... eee es
nemesaunsrijopracnium ruoricatus Allen... 200.2... 60sec. eke eee ele.
menus anerynrus niapyu Aen. 2.0.0.0 .000 5 ce ke we ee ee ees tire
maamenrops veccropi chap Allen... 0... 060. ee ee eb ee eee ee:
ae Aras RRA EG me LU CUM op AEN ys CO TKS, ye ha es ee I a Oe ee
eda OTP UO TM Canteen, ocr ln Sos ore Sia, Seats: aon oo BE SI eS eS
LEGO FLOSS ORIEN CE A ea ee te eg
AOU MME LOU USEC AMNCTY 320.66 50 oie ely oe vow ne wie 6 Ae ee ete es BLS
Re a RUPICROM ODP AULO MIE ee eset A Rech b's Kansas Se oe MOS Chemo an we
minor macrodon Allen... . 0... . RMA AD UM NOTE SMe KTG OC aue 5 Ce teed
NIETO O ECON USS AVICTI . ol ce tesco vee tu kok doa sca py bed ahaa Uae slo aes
LE BO DETALLES A Se oat cect en eee ase MNS nae RP Hea We OER Ce ea
LED LHD 00), ASCII eS RN Te eo a a ace RR Set Ne eT a
MPS TCR MALO US: UTCMSIS NMOM 2.033. 4s0. ee obs lee oe CF ae ee weds tes ee nes
MEV SESCHUGL FONAGIUUS ANEW. 2s oo. hal Sete ols oes Pb hese ces
DDILELP IS CADIS, DAN IST aaa 2h aN Soe ESS artis!

XV"

ERRATA

Page 233, line 16 from top, for Leol. azandicus read Leo l. azandicus,

264, line 2 of table, for dturiensis read iturensis.

287, line 1 from top, for 1625 read 1925.

328, footnote, lines 1 and 2, for synonym read homonym.

359, line 23 from bottom, for LX VII read XLVII.

477, line 14 from bottom, for Fsihego read Fsthego.

477, line 12 from bottom, for Matsche read Matschie.

481, line 19 from top, for Fsihego tturicus read Fsihego ituriensis.

XV1

40 YHomaM ant oT
VddUdTA HIAGA THOU

noitibbs at .odw ,tergolodtiaio bas sergolsmosm tasoimd

betibs bas bstosttb etssy owd-ytudt 10t rodeos as esttub aid ot

IsistsU to museuM oasoremA sdT to ettiomeM bas aitellud ods
.batsoibsb ai e1sqsq evomudieog aid to smulov aidt .yiotetH

as curator

s volume |

thi

SS an ONT aa

BE ie iN

THE AMERICAN MUSEUM OF NaTURAL HISTORY

Vo_ume XLVI], 1922

59.9,33 (67.5)
Article I—THE AMERICAN MUSEUM CONGO EXPEDITION
| COLLECTION OF INSECTIVORA!

By THE LATE J. A. ALLEN?

Puates I To IV, ann 1 Text FIGURE

CONTENTS

PAGE
“CL OTLEVSSELETL DML. 5s cue og ee a rene ae REPS Dh 2

List of Localities, with Names of the Species and Subspecies, and Number
Gyopeemmens taken at each Locality....0.5) a.c2.5 2.00). Fes ee ee 3
New Species and Subspecies, with their Type Localities................. 4
General Summary........ ISS oe SE, BOS ODE ENE Dg RE a NRE 8 YI AEP 4
ie AME TR le 2 oe coe cc Nh on lay ee gh Grd DH NS, Shyer Bb ape SLL MAN Ht Stee eA
MiMCMUBT CRC OTROS (MS oi neta a eek to AT ne Rika t Gi Bois usta ova eels 4
aaa ee a ee ae ote Se eee he ig oie heh es wea taes ney A iuaiee 8
Status of Erinaceus albiventris and E. prunert Wagner................-.. 8
LEE TSS TESTED Loa a laa RR, RS ORES CE 1%
Aa MEMG TUS. NEW SPCClOS io 2 i28 ak) J bo geo da Gh oS Peer ee OM ae we 13
Meet RMN ETCH: SWCCIES & 26 2a Bi <5, 5s duce, Gus) sheesh RS Sinha oleh Ha ee ww Bus eae 15
SaaS UII en SS Oh. tans ake tadueh 4 aces GUERIN wa vaeghen aoe ane 17
TERI E RUE TE EETISCO UCU Ue aN ace cece? os Sec hie wa sooo Gee ee ea eo ar OU wuarelane« 17
UPD DLO? GOP ITED SRI aS ae Ge estes nga nae ane ats a Rn eda 18
“NTE SUEY UNTARELMTI Os CMe RNS BAS eae GA PD Ve eee PE rhe PLE RRL or 18
AOR OC UOMO gc on oN Bare 0s 8 babs vais» nae Bane Leena 18
CIRREGLIRIRE, CLING OT OR N re Oo oe aa ER a EEN aT MST URES Cnt 18
WO ECTOMEMFOCISONT GETUL. 6c ale e's ein ge oe bn ne be BO ys ob) ges cea eee aaah 18
IC TON TMC Oe AOL es Ns Sse isaha A cake. 6) ie gon Sie ah en ee eh aes 18
LAPEER), CITIES, TRE SENG Sak Tee fee Pe mR Bele EM POTEAU) bre ev apmiee Se Tae ato:
ELLE WAS, WEOTET Oe oN RS ee EE AL PO RI ee 19
CDN TERE NTT CAS ie a EDS a a | 2 ee Ett Mi By ced
Pee N ae ROM IO NM ET sR oo Seay gare; © al & au eneyend yesh ot LAR Soke aie 19
BIEL MI OMCTTICOOUCL GLC Me ot ee re 2, a OSE ees eT VAS a 19

= poe Results of The American Museum of Natural History Congo Expedition. Mammalogy,
Oo.

* his paper was in press at the time of Dr. Allen’s death but the final proofs were not seen by him.’
—EpITor.

re, Bulletin American Museum of Natural History [Vol. XLVII

Sylvisorex gemmeus wrene. 60. eee
Sylvisorex Orwunadus. F... occ oA i ne Ue ee eee 20
Scutisorex congicus. 6.5.05 6. oO A a ne ee eee 20
Macroscelididae, .. 0... 62 cle Bee os ee ee ee 20
hynchocyon stuhlmann stuhlmannt.... 06.0... <0... 0
Rhynchocyon stuhlmanni claudi...........-.. 2.2.5 02.
Dentition: of Rhynchocyon. 3.....00.5.6.5 0.0. Sos 1 eee 31
Nastlo fuscipes: on os ge ln kes oe eee eee 37
INTRODUCTION

The collection of Insectivores obtained by the American Museum
Congo Expedition! numbers about 377 specimens, of which 51 represent
the Potamogalidz, 140 the Macroscelidide, 9 the Erinaceide, and 177
the Soricide. In the preparation of this paper the author has had the
efficient codperation of Mr. Herbert Lang, the leader of the American
Museum Congo Expedition. The colored plate (Plate I), illustrating
individual variation in coloration in Rhynchocyon stuhlmanni claudi, is
by Charles R. Knight. The text illustrations are from excellent pen
drawings by Mrs. Ziska.

In working up the material here recorded valuable assistance has
been obtained through material loaned for comparison by the authorities
of the United States National Museum, through the kindness of Mr.
Gerrit 8. Miller, Jr., Curator of Mammals, and from the Museum of
Comparative Zoology at Harvard University, through the kindness of
Director Samuel Henshaw and Dr. Glover M. Allen, Curator of Mam-
mals.

The Soricide of the Congo Expedition were placed for determination
in the hands of Mr. N. Hollister, Assistant Curator of Mammals at
the United States National Museum, in 1916, he having then been for
sometime engaged in a critical study of the African Soricinz in the Na-
tional Museum. His report on the shrews of the Congo Expedition was
published in October 1916.’

1Supplemental Note on Hipposideros lang? Allen (Bull. Amer. Mus. Nat. Hist., XX XVII, pp. 434—-
438, text figs. 4-6. September 29, 1917).

Since the publication of the paper on the Congo Expedition Collection of Bats, I have had an
opportunity to compare, through the kindness of Mr. Gerrit S. Miller, Jr., Curator of Mammalsin the
United States National Museum, three skins and four skulls identified as Hipposideros cyclops (Tem-
minck), from Efulen, Cameroon. While these specimens are not from the type region of cyclops
(Boutry River, Gold Coast), it is interesting to note that they are uniformly and strikingly different
in coloration from the series on which langi was based, indicating at least considerable plasticity in the
cyclops group. Inlangi the whole head is yellowish brown, in strong contrast with the upperparts of
the body, whilein the Efulen specimens it is uniform in color with the back and the upperparts are also
much darker than in langi. The measurements, both external and cranial, indicate slightly larger size
for the Efulen form. Whilelang?, as stated in the original designation, is a member of the cyclops group,

itshould evidently be recognized asa well-marked geographic race, under thedesignation Hipposideros
cyclops langi, and the Efulen specimens as H. cyclops micaceus (de Winton; type locality “Como River,
75 miles from Gaboon’’), with the description of which the Efulen specimens agree and with which the
ldangi series does not agree. Whether or not micaceus is referable in a strict sense to true cyclops I have
not the means at present for determining.

2‘Shrews Collected by the Congo Expedition of the American Museum.’ By N. Hollister. Bull.
Amer. Mus. Nat. Hist., XX XV, pp. 663-680, Pls. vii—x1. October 21, 1916.

1922] Allen, Congo Collection of Insectivora 3

List oF LOCALITIES, WITH NAMES OF THE SPECIES AND SUBSPECIES, AND
NUMBER OF SPECIMENS TakEN AT Eacu LOCALITY

No. of
Localities Species and Subspecies Specimens Totals

Akenge Rhynchocyon stuhlmannii claudi 5
_Avakubi Potamogale velox
_ Rhynchocyon stuhlmanni stuhlmanni
Crocidura nyanse kivu
a S jacksoni denti
bicolor
oritis
latona
Babeyru Crocidura jacksoni denti
Bafwabaka Potamogale velox
x Crocidura jacksoni denti
Scutisorex congicus

ce (a3

‘as ce

its ce

NOR RR KR KR kK OwWNrF ao

igs

Budongo
Forest Rhynchocyon stuhlmanni stuhlmanni
Faradje Potamogale velox
= Nasilio fuscipes
Atelerix faradjius
as = ce langu
Crocidura surure
ne i lutrella
turba nilotica
jacksonz dentr
Sylvisorex gemmeus irene
Gamangui Potamogale velox
a Crocidura nyansxe kivu
Bris a jacksoni denti
Garamba Nasilio fuscipes
Atelerix langi
Crocidura surure
Lubila Crocidura congobelgica
Medje Potamogale velox 30
3 Rhynchocyon stuhlmanni claudr 20
Crocidura nyansxe kivu
oe sd caliginea
yacksoni dente
oritis
latona
ludia
polia
congobelgica
Sylvisorex gemmeus irene
= = oriundus
Scutisorex congicus

i)
LS) RS)
=

ce (as

cc ce

(a9

iss)

SH eee ORE eH oOWwWee o

—

=
I) (0) IRS) SES i I ee)

188

4 Bulletin American Museum of Natural History [Vol. XLVII

No. of
Localities Species and Subspecies Specimens Totals
Nala Rhynchocyon stuhlmanni claudi 1
Crocidura turba nilotica 1
- as jacksoni dent 3 a
Ngayu Crocidura ludia 1 1
Niangara Potamogale velox 3
os Nastlho fuscipes 8
i Crocidura jacksoni dentt 2 13
Niapu Potamogale velox 13
Hy FRhynchocyon stuhlmanni claudi 79 92
Penge Rhynchocyon stuhlmannii stuhlmanni 2 2

New SPECIES AND SUBSPECIES, WITH THEIR TypE LOCALITIES

1. Atelerix faradjius J. A. Allen. Faradje.
2. ‘“langi J. A. Allen. Faradje.
3. Crocidura caliginea Hollister. Medje.
4. ss oritis Hollister... Avakubi.
dD: i latona Hollister... Medje.
6. ludia Hollister.1 Medje.
ie a polia Hollister.1 Medje.
8. i congobelgica Hollister.! Lubila.
9. Sylvisorex oriundus Hollister. Medje.
GENERAL SUMMARY
Species and
Families Genera Subspecies Specimens Localities
Potamogalide 1 1 5l rf
Erinaceidz 1 2 9 2
Soricide 3 15 177 11
Macroscelididze 2 3 140 10
7 21 377
POTAMOGALIDZ

Potamogale velox Du Chaillu

Cynogale velox Du CuatLuv, 1860, Proc. Boston Soc. Nat. Hist., VII (1859-61),
November, pp. 361-363. Ogowe River, French Equatorial Africa. (Potamogale
tentatively proposed on p. 363 in place of Cynogale.)

Mystomys veloz GRay, 1861, Ann. Mag. Nat. Hist., (3) VIII, July, p. 61.

Mythomys veloc Gray, 1861, Proc. Zool. Soc. London, p. 275. Believed to bea
rodent.

Potamogale velox Du CuHaituu, 1874, ‘A Journey to Ashangoland,’ p. 118.
Further notes on the species, in defending himself against erroneous and unkind
criticism.

1Described 1916, Bull. Amer. Mus. Nat. Hist., XXXYV.

1922] Allen, Congo Collection of Insectivora 5

Potamogale velor ALLMAN, 1866, Trans. Zool. Soc. London, VI (1869), pp. 1-16,
Pl. 1 (animal), Pl. 1 (skeleton), text figs. 1-9 (hair, head, ear, feet, anal glands, and
sexual organs). A spirit specimen (dentition not complete, lacking the last molar)
from Old Calabar. |

Potamogale allmani JENTINK, 1895, Notes Leyden Mus., XVI, p. 234.. Based on
Allman’s (as above) detailed account and figures of an immature specimen from Old
Calabar, having only 36 teeth.

Potamogale allmani GranpvIpIER, 1904, Bull. Mus. d’Hist. Nat. Paris, X, p. 51.
Two immature specimens, each with only 36 teeth, provisionally referred to Jentink’s
““espéce incertaine,”’ ‘si son existence réelle était démontrée.”” Of 9 specimens in the
Paris Museum (3 of them without skulls) 6 were yellowish beneath and each of the
skulls, so far as available, had 40 teeth. These were referred to P. velox.

Potamogale velox argens THoMAs, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 470. Two specimens: Medje and Poko, Belgian Congo.

Represented by 51 specimens (with skulls and 7 skeletons), collected
as follows:

Medje, 30 (19 males, 11 females): January 24, 25, March 2-31,
April 10-17, May 9 and 13, August 31, and September 10, 1910; February
27, April 16, June 4, and July 18, 1914.

Bafwabaka, 1: January 9, 1910.

Gamangui, 1: February 20, 1910.

Niangara, 3: November 11-29, 1910; and June 20, 19138.

Faradje, 2: February 21 and May 22, 1911.

Avakubi, 1: December 9, 1913.

Niapu, 13 (10 males, 3 females): November 26—December 31, 1913.

The males number 36, the females 15. The number of fully adult
specimens (of which measurements are given below) is 20 (16 males, 4
females). More than one half are immature, varying from those in
which the third molar, or both the second and third molars, are unde-
veloped (number of teeth 32 or 36) to those with mature dentition (40
teeth).

In respect to seasonal distribution, one or more specimens were
taken in each month of the year except October, but the greater part at
two quite distinct seasons of the year—December and March (November
26—December 31, 13 specimens; March 2-31, 16 specimens). This
would seem to afford opportunity for the study of the influence of season
upon the coloration and character of the pelage, but unfortunately, this
is not the case, since only the Niapu series (taken in December) and a
few others were made up from fresh specimens while the greater part
(including nearly all of the large series from Medje) were not made up
till they were received at the Museum several years later, when it was
found that the fatty matter left on the skins had stained the white under-
parts.

6 Bulletin American Museum of Natural History [Vol. XLVII

Collectors’ measurements of 7 adult males and 3 adult females from

Medyje:
Total Length Head and Body Tail Vertebree Hind Foot Ear

g 575 (555-610) 310 (298-325) 267 (250-290) 43.0 (41-46) 21.5 (21-28)
g 603 (600-610) 323 (310-339) 284 (280-290) 41.7 (41-438) 21.5 (21-22)

Skull, same specimens, condyloincisive length: <7, 63.8 (61.1-65.9) ;
©, 64 (63.6-64.3).

Collectors’ measurements of 5 adult males and 1 adult female from

Niapu:

Total Length Head and Body Tail Vertebre Hind Foot Ear
o 586 (542-610) 329 (297-347) 260 (245-273) 45 (43-48) 22 (22-22)
Q@ 585 330 ABE 4] 21

Skull, same specimens, condyloincisive length: 7; 65.5 (63.8-66.5) ;
OF Gare: ,

Collectors’ measurements of 4 adult males from other localities (Baf-
wabaka 1, Gamangui 1, Niangara 2):

Total Length Head and Body Tail Vertebre Hind Foot Ear

565 (550-583) 294 (279-3812) 268 (260-272) 48.5 (43-44) 21.3 (20-22)

Skull, same specimens, condyloincisive length: 64.8 (62.7—-66.7).

The above statistics indicate that there is no distinctive sexual
difference in size. The smallest skulls in each series are, as a rule, the
youngest, or skulls with full mature dentition but in which the teeth are
wholly unworn and the cranial sutures still distinct. There is no skull
young enough to show the milk dentition. In several skulls in which the
first molars are fully developed, the condyloincisive length is 50-51 mm.;
in those in which the second molars are fully up but the third molar is
still enclosed in the gum it ranges from 53-55 mm.; during the develop-
ment of the third molar the skull length increases to about 60-61. 5.

Potamogale velox argens was based on two specimens, one from Medje
and one from Poko. As no type locality was definitely indicated, Medje,
the first locality mentioned in the description, is here designated as the
type locality. Hence the present series of 31 specimens from Medje are
topotypes. Poko and Niapu are both near Medje. The Niapu series of
13 specimens is in fine condition, the underparts being unstained, and
agrees in a general way with the brief description of argens; it shows,
however, that the white area of the underside varies in extension upward
on the sides and that the development of brown-tipped hairs along its
upper border is also a variable feature. The fore limbs are sometimes
‘‘almost wholly in the whitish area’’ and sometimes wholly brown above
in specimens from the same locality and collected on the same day, show-

1922] Allen, Congo Collection of Insectivora é

ing this alleged distinction to be subject to a wide range of individuality.
Specimens taken at approximately the same date vary greatly in the
condition of the pelage in respect to wear, but, on the whole, December
specimens, taken near the close of the dry season, appear more worn than
those taken in March—May, the rainy period. But doubtless the season
of moult varies in different individuals and, like the birth of the young,
may extend over a considerable part of the year.

Only two forms of Potamogale, in addition to the original P. velox
Du Chaillu, appear to have been distinguished. These are P. allmani
Jentink (1895), based on Allman’s detailed description (loc. cit.) of a
specimen preserved in spirits from Old Calabar, published some twenty-
six years before. The second, P. velox argens Thomas, was added in

- December 1915, on the basis of two specimens from the Upper Congo.

The large series of specimens collected by the American Museum Congo
Expedition demonstrates that the characters relied upon for the discrim-
ination of these two forms are without value and, for this reason, are
subjected to comment.

As shown in the collectors’ field notes on this species the genus
Potamogale has a wide geographic range, and hence might be expected
to have developed local phases. It is not the purpose of these remarks
to discredit such a reasonable probability but merely to show that the
evidence presented for the two forms above cited is far from adequate.
Unfortunately, little material is available for direct comparison with
that from the Upper Congo region, but the latter emphatically shows the
trivial nature of the distinctions offered by their describers for the recog-
‘nition of allmani and argens. Reference has already been made (p.
5) to the stained condition of the underparts due to treatment of the
skins before they were made up. Apropos of this, and in response to my
inquiries, the collectors have informed me that ‘‘all living or freshly
killed specimens they saw had pure white, lustrous fur on the under
side, if not soiled by the reddish clay of these regions”; and they add
that “some of their own skins from the same places, when unpacked,
were yellowish, due to a difference in the method of preservation.”’
They also state that they ‘‘noticed that in many old, flat skins or por-
tions of them from the same localities, which they saw in the possession
of Europeans and natives, the originally white area was always yellowish
or brown.”’ It is also well known that in museum specimens the white
underside of mammals long stored are apt to turn yellowish from fatty
matter retained in the skin, or from other causes, and therefore are un-
satisfactory as standards of comparison with freshly collected material.

8 Bulletin American Museum of Natural History [Vol. XLVIT

While Potamogale is a rather common animal in its native haunts, it is
one of the rarest in collections and, when present, is doubtless not safely
comparable with freshly killed specimens.

Potamogale allmani was proposed on the basis of two (in part hypo-
thetical) distinctions: (1) the presence of 36 teeth (owing to the im-
maturity of the specimen) instead of 40; (2) the ‘‘brownish yellow”
instead of white underparts, due to discoloration by the preservative.
P. velox argens was described from two apparently normal specimens in
which the white of the underparts reached “‘higher up on the sides”’ of
the body and on to the under surface of the basal portion of the tail than
in P. velox, which features a large amount of topotype material shows to
be inconstant and merely individual. Consequently allmani and argens .
cannot be considered as entitled to recognition.’

It is hardly necessary to add that many forms, species as well as
subspecies, have a similarly unsatisfactory basis, as they rest on slight
differences shown by single specimens, or on characters of trivial import-
ance. Their confirmation, it is obvious, rests on a comparison of ade-
quate series of topotype material with similar series of their near allies;
and the author who would discard them without such resources would
take great risks, notwithstanding his strong conviction that the forms in
question are merely names.

ERINACEIDE ;
Status oF Hrinaceus albiventris AND E. prunert WAGNER

The Erinaceide are represented in the Lang-Chapin Congo Col-
lection by nine specimens (skins with skulls), all from Faradje except a
third-grown female from Garamba, a nearby locality. They comprise
three adult females and six young, from one-third to one-half grown, and
unquestionably represent two distinct. species, differing in important
cranial characters and in external features. Both belong to the section of
the family in which the hind feet are four-toed. Owing, however, to the
unsatisfactory original descriptions of the first-named members of this
group, and to lack of proper material for direct comparison with the
Lang-Chapin specimens, their determination has been difficult. Thanks
to the authorities of the Museum of Comparative Zoédlogy at Harvard
University and of the United States National Museum at Washington,
I have in hand 14 additional specimens of the group with 4-toed hind

1In respect to the status of P. allmant, cf. Leche, who in 1907 Pegler XX, Heft 49, pp. 6 and
129, footnote 1) regarded it as only ‘‘ein jugendliches Individuum”’ of P. velox.
2S8ee above (p. 5) the citation of Grandidier’s paper on the Paris Museum series of Potamogale

1922] Allen, Congo Collection of Insectivora 9

feet, making 23 in all, representing five easily recognizable forms. While
their relationships inter se are obvious, the names properly applicable to
the two forms from Faradje have raised a serious question of nomen-
clature. One of them should apparently be referred to EH. albiventris
Wagner, as that name has of late been employed, but which of them
should be so recognized is indeterminable. This raises the further and
more fundamental question of the availability of this name, considered
with relation to its origin and history.

As is well known, Wagner, in 1841, described as new two species of
Erinaceus (albiventris and pruneri) on consecutive pages of the same
work,! for neither of which was given a definite type locality. Hrinaceus
albiventris, the first in sequence of the two species, was based on a single
specimen obtained from a dealer, who stated that it was found in a, col-
lection from India (‘‘befand er sich unter einer Sendung indischer
Thiere’’), Wagner himself saying: ‘‘ Die Heimath kann ich nicht genau
bestimmen.”’ The original description of the species was inadequate,
merely indicating that it had, like many other species of Erinaceus now
known, white underparts, parti-colored spines, and other features of no
distinctive significance. In later references? to the species he stated that
the hind feet have only four toes. This fixed the ‘‘Heimath” as Africa,
inasmuch as no species of this genus having 4-toed hind feet are known to
occur elsewhere. Fortunately, the type remained available for examina-
tion by later investigators, confirming its African origin. Erinaceus
albwentris Wagner thus became a “blanket name”’ for all the African
. species of Erinaceus with 4-toed hind feet. Various forms of the group
later became segregated, one after another, under distinctive names as
species, and the name albiventris, by some authors, was POR Cet (appar-
ently rather informally) to a Sudan form.?

Erinaceus prunert Wagner, synchronous in publication with his E.
albwentris, was based on specimens received from Dr. Franz Pruner,
from a locality not definitely stated in the original description, nor in
Wagener’s later references to the species,* where he gives its distribution
as ““Sennaar, nach Sundevall auch am Senegal.”’ It is to be noted that he
synonymized (in 1842 and 1855) E. heterodactylus Sundevall, based on
specimens from the Bahr el Abiad (White Nile), Sennaar, with his ZL.

~ St

11841, ‘Schreber’s Siugthiere,’ Suppl., II, pp. 22 and 23.

seer Wiegmann’s Archiy fiir Naturg., 1X, 2 Bd., p. 27; 1855, ‘Schreber’s Siugthiere,’ Suppl., V,
p.5

3Thomas and Wroughton, in describing their Erinaceus spiculus, from near Lake Chad, in 1907
(Ann. Mag. Nat. Hist., (7) Nie p. 371), made comparison with ‘‘the Soudanese BUCS z
and further state: ‘‘ The nearest neighbors of spiculus are albiventris, Wagner, from the Scucan and
Adansoni, Rochebrune, from Senegal.’’

41855, ‘Schreber’s Saugthiere,’ Supp!., V, p 587.

10 Bulletin American Museum oj Natural History [Vol. XLVII

pruneri, a course followed by apparently nearly all subsequent authors.
But not by all, since von Heuglin! in 1867 gave a list of the species of
Erinaceus occurring in ‘‘ Nordost Afrika” in which he included: ‘ Erina-
ceus prunert Wagner. Aus dem Sennaar.’”’ And Fitzinger? recognized it
as not only distinct from albzventris but as the type and only species of
his genus Peroéchinus. In the original description of prunerz it is stated
merely that the specimens on which it was founded came in a collection
of mammals sent by Dr. Pruner from ‘ Kairo.”’ It is known, however,
that Dr. Pruner visited the Upper Nile region and there collected speci-
mens of hedgehogs that were sent to Wagner,? among them those on
which prunerz was originally based.

It may be noted further that Sundevall, about the same time (see
below, p. 12), described his Erinaceus heterodactylus,* a species having
4-toed hind feet, based on specimens collected by Dr. Hedenborg in
“Sennaar,”’ and that this species has always been synonymized with £.
prunert by subsequent writers, both forms coming from ‘“‘Sennaar.”’
As both have been referred by most authors to E. albiventris, it may be
that this fact has had an influence in the recognition of Sudan as the
type region of albiventris (or, more definitely, Kordofan, in the case of
prunert).

To follow further the history of EF. albiventris, from a geographic
point of view, Fitzinger, in 1867,° gave its Vaterland as “‘nicht mit
Sicherheit bekannt, wahrscheinlich aber Ost-Indien,” and that of
pruneri as Kordofan. Dobson, in 1882,° gave the range of albiventris as
“Northern Tropical Africa (Senegambia, Sennaar),’’ and Anderson in
1895,’ as extending from ‘‘Senegambia across Central Africa, south-
wards to Uganda and northwards to Somaliland.”’ In 1902,’ he stated:
“The specimen upon which Wagner founded this species [Hrinaceus
albiventris| came, in all probability, from Senegambia,’? and adds:
“The Nile Valley and East African specimens . . . may be more

1‘Beitr. zur Fauna der Siuget. N. O. Afrika,’ p. 22.

21867, Sitzungsb. math. nat. Cl. Akad. Wiss. Wace LVI, p. 126.

3Cf. Anderson’s ‘Mammals of Egypt,’ 1902, p. 162.

41841, Sven. Vet. Akad. Handl. Stockholm, (1842,) p. 227.

5Sitzungsb. math. nat. Cl. Akad. Wiss. Wien, LVI, p. 856.

6* Monograph of the Insectivora,’ p. 11.

7Proc. Zod]. Soe. London, 1895, p. 420. Andersonincluded under Erinaceus albiventris E. prunert
Waener, E. heterodactylus Sundevall, HE. adansoni Rochebrune. He added: ‘This species [albiventris]
has been obtained at the following localities: Senegal; Saint Louis; Cape Verd; Joal; MacCarthy’s
Island, River Gambia [collectively =range of EH. adansoni]; Accra, Fantee; Porto Seguro, Togo;
Gaboon; Kitui, Ukamba [type locality of E. hindei Thomas, 1907]; Tabora; Kasé; Kilima Njaro;
Wakilomi, District of Maka; Central Somaliland; Sennaar [E. heterodactylus]; Kordofan [E. pruneri];
and region of Upper Nile.’

8‘ Mammals of Egypt,’ p. 164.

*This statement, daabtless was based on his personal examination of the ty pe in the Munich Mu-
seum, as he statesin another connection (1895, Proc. Zo6]. Soc. London, p. 414): ‘‘I may mention that I
have examined all the Hedgehogs preserved in the Museums of Paris, Frankfort on the Main, Munich,
Berlin, and London, and . . . some of the specimens described by Fitzinger in the Vienna Museum.”

1922] Allen, Congo Collection of Insectiora 11

definitely registered as Erinaceus albiventris subsp. prunert.” Again
he says, later on the same page, referring to albzventris: ‘‘This species is
found to the south of Khartum,” and ‘“‘ranges into Somaliland and as
far south as Kilima-nyaro.”’ Within this regicn, since 1902, two forms
(E#. hindet Thomas and E. albiventris sotike Heller) in addition to pruneri
have been recognized,! and two more are added in the present paper,
both from Faradje, northeastern Belgian Congo.

As stated above (p. 9, footnote), Thomas and Wroughton in 1907,
in describing their Erinaceus spiculus, referred to Sudanese specimens
as typical of E. albiventris. In view of the complications of the case, it
seems to me preferable to place Erinaceus albiventris permanently in the
list of unidentifiable species, it having no type locality and being speci-
fically unidentifiable from the original description, although the type
appears to have been preserved in the Munich Museum.” Senegal (or
Senegambia) and Sudan (or Sennaar), the rival suggested type regions,
are far apart, with EL. adansoni representing the former and EF. prunerz
the latter as well established species. Under this ruling the two forms
from Faradje are described as new. |

ATELERIX POMEL

Since the foregoing was prepared for the press a paper by Oldfield
Thomas, on ‘The Generic Divisions of the Hedgehogs’ (1918, Ann. Mag.
Nat. Hist., (9) I, February, pp. 1938-196) has appeared, respecting which
a few notes are here appended. The old genus Erinaceus is divided by
Thomas into five genera, which, with their designated genotypes, are as
follows: -

1. Erinaceus Linné, 1758. Genotype, E. europxus Linné.

2. Athechinus, new genus. Genotype, LE. algirus Duvernoy and
Lereboullet.

3. Atelerix Pomel, 1848. Genotype, E. albiventris Wagner.

4. Hemiechinus Fitzinger, 1866. Genotype, E. platyotis Sundevall.

5. Parechinus Trouessart, 1879. Genotype, HE. micropus Blyth.

Although each of these groups is represented in Africa, only
Atelerix and Athechinus come geographically within the scope of the
present paper. Atelerzx was proposed by Pomel (1848) as a subgenus of
Erinaceus, with the statement ‘‘4 dactylus”’ as the entire diagnosis. No
species was referred to it, and no geographic range was indicated for the

1Since this was written Thomas has added a third from Kilimanjaro as Atelerix kilimanus (1918,
Ann. Mag. Nat. Hist., (9) I, March, p.232.

2As stated above (p. 10, footnote), the type was probably critically studied by Anderson prior to
1895, together with the type of prunert, leading to his assignment of the type locality of albiventris to
Senegambia, and to his later recognition of pruneri as an eastern subspecies of albiventris.

12 Bulletin American Museum of Natural History i[Vol. XLVII

group. Neither is it indicated whether ‘4 dactylus”’ refers to the hind
feet or to the fore feet, or to all the feet. It happens, however, that only
one species of hedgehog had at that time been characterized as 4-dactylus
in the original description of the species. This was Erinaceus hetero-
dactylus Sundevall (1841, Sven. Vet. Akad. Handl. Stockholm, p. 227),
which is characterized as, among other distinctions, ‘‘Pedibus posticis 4
dactylis,’”’ which is doubtless the original source of Pomel’s ‘4 dactylus.”’
At about the same date (1841) Wagner described Erinaceus albiventris
and H. prunerz on consecutive pages of the same work, without specify-
ing this character for either species. The first of these (EZ. albiventris) I
consider specifically unidentifiable, for reasons already given in the
present paper. This is the species now designated by Thomas as the
type of Ateleriz. Wagner, two years later, in his ‘Bericht tiber die
Naturgeschichte der Siugthiere wahrend des Jahres 1842’ (1843, Arch.
fir Naturg., Bd. 2, p. 27), claimed priority for his prunerz over hetero-
dactylus Sundevall, to which he referred the latter as a synonym. He
says he received a separate of Sundevall’s paper from the editor of the
Archiv, and that the volume in which it was printed was issued later, but,
as he fails to state when Sundevall’s paper was received, or what date it
bore, we are left in doubt as to which paper has priority of publication,
the date of his own publication being ‘15. Mai 1841.”

In his comment on Sundevall’s paper he says that “‘ LE. heterodactylus
Sund. mit meinem E. Pruneri identisch ist; auch der hintere Daumen
geht diesem wie jenem ganz ab.”’ He says further that he had assumed
the absence of the hallux in E. pruneri and E. albiventris to be the result
of an injury and for that reason did not mention it; but, inasmuch as
Sundevall had found the same suppression in his EF. heterodactylus, he
now considered it an important character for his EH. prunert and H#.
albiventris, to be included in the diagnosis. It is accordingly so included
in his later revision of the hedgehogs (1855, ‘Schreber’s Saugt.,’ V, p. 587).

The question of what name the genotype of Atelerzz should bear is
thus somewhat complicated, depending upon priority of publication of
the names 'E. heterodactylus Sundevall, under which the expression ‘4
dactylus”’ (the sole diagnosis of Atelerix) was first employed for a hedge-
hog, and which was first recognized as a character of EH. pruneri some
two years later. In any case, by the consensus of authorities both names
refer to the same species. Furthermore, Peroéchinus Fitzinger (1866),
without diagnosis, included only HE. prunerz (with E. heterodactylus
Sundevall as synonym), which is, therefore, the genotype of Peroéchinus.
As Peroéchinus is a substitute name for, or at all events a pure synonym

1922] Allen, Congo Collection of Insectivora 13

of, Atelerix, it thus determines under the peculiar conditions of the
case the genotype of the latter as E. pruneri. (Cf. ‘Internat. Code Zool.
Nomen.,’ Art. 30, II, f.)

In Thomas’s synopsis of the hedgehogs, the sole distinctive character
of Atelerix is: ‘‘Hallux absent;”’ and, so far as I can find, this is the only
distinction between Ateleriz and his new genus 4thechinus, defined as:
“Coronal parting broad, conspicuous. Posterior palatal shelf broad.
Third incisor two-rooted.”’ The last two characters, in comparison with
Erinaceus (as restricted by Thomas), are both present in Ateleriz;
the first is of less importance, depending upon the stress to be laid upon
the words “‘broad, conspicuous,” since in Atelerix there is a distinct
coronal parting, although less developed than in Erinaceus europxus
and its near allies.

As shown below (p. 17), the absence of the hallux is not constant,
and therefore not an important character, since in different individuals
of the same litter of young it may be present or absent, although absent
as a rule in a number of forms of the prunerz (heterodactylus ?)-adansoni
group, which is distributed over a wide geographical area. I agree with
Thomas that both Atelerix and Athechinus are separable from Erinaceus,
sensu stricto, but collectively rather than as two generic groups, for which
the rule of priority demands the earlier name, Ateleriz.

The forms referred to Atelerix by Thomas are:

1. albiventris =Erinaceus albiventris Wagner, 1841.

2. adansoni =F. adansoni Rochebrune, 1882.

3. hinder =F. hindet Thomas, 1910.

4. spiculus _=E. spiculus Thomas and Wroughton, 1907.
5. spinifex = Atelerix spinifex Thomas, March, 1918.

6. kilimanus =A. kilimanus Thomas, March, 1918.

To which may be added:

7. hinder sotike =E. sotike Heller, 1910.
8. faradjius =A. faradjius (described below).
9. langi =A. langi (described below).
And pruneri=Erinaceus pruneri Wagner, 1841 (=?E. heterodactylus
Sundevall, 1841), in place of ““albiventris’’ as No. 1 of the above list,
and also as type of Ateleriz in place of albiventris.

Atelerix faradjius, new species
Type, No. 51006, 9 ad., Faradje, northeastern Belgian Congo, July 7, 1911;
Herbert Lang and James P. Chapin. American Museum Congo Expedition. Orig.
No. 1660. Topotype ( 9 very old), No. 51007.

Represented by two adult females from Faradje, of the so-called “albiventris’’

type.

14 Bulletin American Museum of Natural History (Vol. XLVII

General coloration of the upperparts strongly yellowish white superficially, the
broad light tips of the spines being of this color and nearly concealing the dark sub-
terminal zone. Head in front of eyes, including sides of nose, dull tawny-brown;
also ears and feet the same in general effect; a broad frontal band, cheeks, sides of
neck, sides of shoulders and forearms, thighs and hind legs, rump and whole under-
parts uniform dull yellowish white (possibly white slightly stained yellowish); upper
surface of fore feet slightly clothed with yellowish-white hairs, hind feet more heavily
clothed with longer yellowish-white hairs, through which the pale tawny color of the
skin determines the general effect; tail similar in coloration to the feet. Spines
broadly tipped (for about 4-5 mm.) with yellowish white (without darker tips);
subapical band (about 5 mm.) dark tawny-brown, passing proximally into dull yel-
lowish white on the basal half. Longest head spines about 17 mm. in length, body
spines about 15 mm.

Collectors’ measurements: total length (type), 249 mm.; head and body, 230;
tail, 19; hind foot, 29; ear, 30. Topotype (very old female with greatly worn teeth) :
total length, 205; head and body, 180; tail, 25; hind foot, 26; ear, 30.

Skull measurements: condyloincisive length, (type) 45.1, (topotype) 43.6;
length of nasals, 16.5, 15; palatal length (to front of premaxille) 25.4, 24.7; zygo-
matic breadth, 27.7, 26.3; interorbital breadth, 11.8, 11.5; breadth of braincase,
19.5, 19.9; postglenoid breadth, 22, 20; mastoid breadth, 15.5, 16.5; palatal breadth
(outside to outside of m!), 17.6, 16.8; breadth of rostrum at base of front incisor, 6, 5.4;
breadth of palate at ridge behind m®, 9.7, 8.5; tip to tip of alisphenoid processes,
11.2, 11.3; tip to tip of pterygoids, 6.1, 6.7; length of mesopterygoid fossa, 10.7,
10.2; breadth between pterygoids, 2.8, 2.7; length of upper toothrow (i!—m*), 21.5,
21.5; upper molars, 8.1, 8.1; lower toothrow (to tip of i’), 9.7, 8.2; lower molars, 9.9,
9.8; length of mandible (front of symphysis to posterior border of condyle), 34.5,
34.2; depth, angle to coronoid, 17.7, 16.7.

The skull is large and heavy; the nasals are long and narrow, the premaxillze
greatly extended posteriorly, meeting the frontals and excluding contact of the maxil-
le with the nasals; zygomatic arches narrow as in A. pruneri; mesopterygoid fossa
very broad, the pterygoids and alisphenoids heavily developed and widespreading as
in A. hindei (the reverse of what is seen in A. pruneri'!); dentition heavy, as in A.
langi and A. hindet.

The pattern of coloration is as in A. prunerz, differing from that of
langi and hindez in having the space below the eye white instead of black~
ish. The spines are as in pruneri—short and fine instead of long and
coarse, and those of the frontal border not conspicuously lengthened as
in the hinder group. The general coloration of both spine-tips and hair
is more yellowish and less clear white than in prunerz; the nose and basal
color of the feet and ears is tawny instead of blackish as in prunerz and
in hindei. This however may be subject to considerable variation through
seasonal and other conditions.

1The specimen of A. pruneri here employed in comparison is No. 14446, Mus. Comp. Zo6l., a young
adult male (teeth unworn), collected at Fazogli, Blue Nile, by Dr. G. M. Allen and recorded by him
(Bull. Mus. Comp. Zo6l, LVITI, p. 342, July, 1914) as Hrinaceus albiventris pruneri.

1922] Allen, Congo Collection of Insectivorc 1d

Atelerix langi, new species

Type, No. 51000, 9 ad., Faradje, northeastern Belgian Congo, March 22, 1911;
Herbert Lang and James P. Chapin. American Museum Congo Expedition. Orig.
No. 1544.

Represented by 7 specimens, the type, an old female, and her litter of five young
(3 males and 2 females), about one-third grown, taken March 22, 1911, at Faradje,
and another third-grown young collected at Garamba, May 1, 1912.

A dark-colored species, allied to Erinaceus hindei Thomas of British East
Africa.

Typre.—Upperparts dark brown, the spines over the greater part of the back
uniform blackish brown from base nearly to tip, the extreme tips tending to lighter
brown or even whitish; front of head, flanks and posterior margin of back lighter
than the mid-dorsal area, the spines distinctly whitish-tipped, especially on the lower
back where all are conspicuously whitish terminally. Ventral surface white, the
white area extending along sides of body, shoulders and forearms, and joining the
broad white frontal band between the eyes and base of the ears. A narrow line of
dusky brown borders the white band in front, broadening laterally to include the
cheeks below the eye and extending forward to the naked portion of the face, which,
with the chin, is also dark brown. Upper surface of fore and hind feet dark brown,
but much lighter than the cheeks. Ears and tail dull brown, the former nearly naked.

Younc.—The five third-grown young differ uniformly from the adult type
specimen in the dark markings of the face being more intensely black, in vivid contrast
with the clear white frontal band. The upper surface of the feet is also deep blackish
brown, as is also the tip of the inconspicuous tail. The spines of the dorsal area are
all conspicuously and uniformly tipped with white, through which the blackish brown
proximal portion of the spines is more or less visible. The young specimens have a
tendency to a narrow blackish median area on the posterior part of the ventral surface,
in some of them strongly developed. They agree strictly with the mother in the
color pattern, but have the black on face and feet more intense and more sharply
defined, and the white or whitish tips to the spines longer. The slightly younger
specimen from Garamba is indistinguishable from the Faradje specimens in coloration
and details of structure.

Collectors’ measurements of the type: total length, 195 mm.; head and body,
175; tail, 20; hind foot, 28; ear, 21.

Skull: condyloincisive length, 43.3; length of nasals, 15.4; palatal length (to
front of premaxillz), 24.3; zygomatic breadth, 29.4; interorbital constriction, 11.7;
breadth of braincase, 18.3; postglenoid breadth, 21.7; mastoid breadth, 14.7;
palatal breadth (outside to outside of m!), 17.9; breadth of rostrum at 1!, 6.6; breadth
of palate at ridge behind m®, 8.2; tip to tip of alisphenoid processes, 9.2; tip to tip
of pterygoids, 5.4; length of mesopterygoid fossa, 10.5; breadth of fossa between
pterygoids, 3; length of upper toothrow (i'—m’), 21.3; upper molars, 9; lower tooth-
row (tip of i;-m3), 20.2; lower molars, 10.2; length of mandible (front of symphysis
to posterior border of condyle), 33.5; angle to condyle, 17.5.

In pattern of coloration A. langi agrees with A. hindez, in both the
dark color of the face extending over the cheeks, which are white in
faradjius and pruneri; indeed, the series of young specimens of langi
are almost indistinguishable in external features from a corresponding

16 Bulletin American Museum of Natural History [Vol. XLVII

series of young sotike (a slightly differential form of hindez). The dorsal
coloration in both is superficially dark brown in general effect but the
single adult of Jang: is much darker than any of the four adults of hindez
available for comparison,! while the white tipping of the spines is con-
spicuous and uniform in hindez and nearly absent in langi. The spines in
lang: are blackish brown from tip to base, lacking the light median band
present in the hindez group. The interaural spines in both are much
longer than those of the body, forming a decidedly lengthened frontal
crest, absent in the prunerz (‘“albiventris’’) group.

_ The type skull agrees in general dimensions with those given for the
type of hindez, but differs from it in the nasals being much longer; the
short nasal border of the premaxille, with a naso-maxillary junction as
long or longer than the nasal contact with the premaxille—dquite the
reverse of the conditions in hindez, in which the premaxille are “slanted
backwards, touching the tips of the frontal processes and shutting off
the maxille from the nasals.’”’ The postpalatal region is also much nar-
rower, the pterygoid and alisphenoid processes weaker and much less
everted, thus giving to this region a quite different aspect. All of the 6
young skulls (of which the type is the mother of 5 of them) agree with the
type skull in the short naso-premaxillary suture and the long naso-
maxillary suture, and the narrow postpalatal region and weak develop-
ment of its processes.”

Of 12 skulls of the hzndez group (5 of hindez and 7 of sotike [E. albi-
ventris sotike Heller], the latter all from the Guaso Nyiro River) all but
one have the nasal border of the premaxille extended posteriorly
(‘slanted backwards’’), and in all but two they nearly or quite reach the
frontal processes, the maxille not reaching the nasals or barely touching
them for usually less than a millimetre.

The skull of the type of hindez (a female) appears to have been ex-
ceptionally large (‘‘greatest length 44; zygomatic breadth 30 mm.”),
none of the four adult male skulls before me exceeding a total length of
43 mm., with an average of 42.1, and a maximum zygomatic breadth of
27.6, with an average of 26.7, although the teeth are worn and one
(total skull length 41.7) is very old. The author’s suggestion that when

1The specimens of hindei available for comparison are: Nos. 16096 Mus. Comp. Zoél., subadult 0,
Upper Ura River; 16097 Mus. Comp. Zo6l.; ad. @ (skin only); No. 161699 Nat. Mus., ad. o’, Kapiti
Plains; Nos. 164022 and 164023 Nat. Mus., both ad. 9, Ulucania Hills; No. 182652 Nat. Mus., very
old &@ (teeth greatly worn), Lololokwi. All the localities are in British East Africa, not far from the
type locality of hindei (Kitui, about 75 miles southeast of Mt. Kenia). The Mus. Comp. Zo6l. speci-
mens (both ex Wulsin Coll.) are labeled Erinaceus hindei; the Nat. Mus. specimens, Hrinaceus albi-
ventris hindet.

‘ 2In respect to this latter feature comparison is made with skulls of hindei from the type region of
*he species.

1922] Allen, Congo Collection of Insectivora iw

males are available for examination they would prove to be larger than
the type is thus not confirmed by the present material.

The hind feet in the type of A. lang: show no vestige of a hallux.
The slight taxonomic significance of its presence or absence in this genus
is well indicated by the series of 6 young specimens of which the type of
langi is the mother of 5. Of the 6 young ones 3 have a vestigial hallux and
the other 3 are without it. It is also much more developed in one of the
three in which it is present than in the other two. It is also present in one
of the 3 young sotzke specimens (No. 181441 Nat. Mus.), and absent in
the other two and in 12 adults of the hindez-sotike series.

SORICIDZA

As stated above (p. 2), the shrews of the Congo Expedition were
early assigned to Mr. Hollister, of the United States National Museum,
for determination, and his report on them was published in this Bulletin
in October, 1916.1! The following statement in respect to the extent and
character of the collection is made in the introduction to his paper:

The shrews collected by Herbert Lang and James P. Chapin on the American
Museum Congo Expedition number 183 specimens, of 15 species and 3 genera.
Almost one half of the species are new. This is not altogether surprising when it
is considered how few shrews have been described from the Congo as compared
with other parts of Africa. It nevertheless seems remarkable that five of these new
species should be members of the small group of “naked-tailed”’ Crocidura of which
only about ten forms were heretofore known. Five forms of Crocidura which have
been recorded from the general region are not represented in this collection. These
are Crocidura turba turba Dollman, C. t. tarella Dollman, C. poensis attila Dollman,
C. boydi Dollman, and C. nigrofusca Matschie. Races of C. hildegardex and C.
fumosa, as well as representatives of several west coast species also might reasonably
be expected.

In order to complete the record of the Congo collection of insecti-
vores, the shrews obtained are here listed, as determined by Mr. Hol-
lister.

Crocidura nyanse kivu Osgood
Plate IV

Crocidura nyanse kivu HOLvisTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV,
p. 663, Pl. x, fig. 1 (animal).

Crocidura nyanse kivu ALLEN, 1917, Bull. Amer. Mus. Nat. Hist., XX XVII,
pp. 769-774, figs. 1 and 2 (skull), figs. 5-8 (skeleton), Pl. xc1r (animal, from photo-
graph). Skull, skeleton, and external appearance, in comparison with Scutisorex
congicus.

Specimens, 20: Avakubi, 3 (1 alcoholic); Gamangui, 1; Medje, 16.

1‘Shrews Collected by the Congo Expedition of the American Museum.’ By N. Hollister, Bull.
Amer. Mus. Nat. Hist., XX XV, pp. 663-680, Pls. vii—x1. October 21, 1916.

18 Bulletin American Museum of Natural History [Vol. XLVII

Crocidura surure Heller
Crocidura surure HouuistER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 664.
Specimens, 2: Faradje, 1 (skin and skull); Garamba, 1 (alcoholic).

Crocidura lutrella Heller
Crocidura lutrella HoLutstErR, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 664.
Specimens, 1: Faradje (skin only).

Crocidura turba nilotica Heller

Crocidura turba nilotica HOLusTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV.
p. 664.
ye = =Crocidura turba nilotica ALLEN, 1917, Bull. Amer. Mus. Nat. Hist., XX XVII,
p. 784, Pls. Lxxx1x and xc (skiagraphs of skeleton, in comparison with skeleton of
Scutisorex congicus).

Specimens, 4: Faradje, 3 (2 Ac: Nala, 1 (alcoholic).

Crocidura caliginea Hollister

Crocidura caliginea HouutstER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p.
664, Pl. vu, fig. 1 and Pl. vim, figs. 1, 1a (skull).

“Type, No. 48555, Amer. Mus. Nat. Hist., skin and skull of adult 2
(teeth moderately worn and basal suture closed) collected at Medje,
Belgian Congo, July 8, 1914, by Herbert Lang and James P. Chapin.
Orig. No. 2451.”

Crocidura jacksoni denti Dollman
Plate II, Figure 1
Crocidura jacksoni denti HouLuisTER, 1916, Bull. Amer. Mus. Nat. Hist., MXXKV,
10, OO, Telos nes 2
Specimens, 71: Avakubi, 6; Babeyru, 1 (alcoholic); Bafwabaka, 1;
Faradje, 5; Gamangui, 2; Medje, 51; Nala, 3 (alcoholic); Niangara, 2.

Crocidura bicolor Bocage
Crocidura bicolor HouuisTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 666.

Specimens, 1: Avakubi (alcoholic).

Crocidura oritis Hollister
Crocidura oritis HOLLISTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 666,
Pl. vu, fig. 2 and Pl. vim, figs. 2, 2a (skull).

“Type, No. 48510, Amer. Mus. Nat. Hist., skin and skull of adult <7
(basal suture closed; teeth moderately worn) collected at Avakubi, Ituri
River, Belgian Congo, July 6, 1914, by Herbert Lang and James P.
Chapin. Orig. No. 2530.”

1922] Allen, Congo Collection of Insectivora 19

This species is based on five specimens, four from Medje and one
(the type) from Avakubi.

Crocidura latona Hollister
Crocidura latona HouuistTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 667,
Pl. vu, fig. 3 and PI. vit, figs. 3, 3a (skull).
“Type, No. 48610, Amer. Mus. Nat. Hist., skin and skull of adult 7
(basal suture closed; teeth moderately worn) collected at Medje, Bel-
gian Congo, March 17, 1910, by Herbert Lang and James P. Chapin.

Ones iNo- 773.”
_ Besides the type there is a single skin without skull, from Avakubi.

Crocidura ludia Hollister

Crocidura ludia HouuisTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 668,
Pl. vu, fig. 4 and PI. rx, figs. 1, 1a (skull).

“Type, No. 48566, Amer. Mus. Nat. Hist., skin and skull of adult o
(teeth slightly worn and basal suture not closed) collected at Medje,
Belgian Congo, May 16, 1914, by Herbert Lang and James P. Chapin.
Orig. No. 2366.”

There are three specimens of this species in the collection, two
from Medje (one the type) and.one from Ngayu.

Crocidura polia Hollister
Crocidura polia HouuistTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p. 669,
Pl. vii, fig. 5 and PI. 1x, figs. 2, 2a (skull).

“Type, No. 48559, Amer. Mus. Nat. Hist., skin and skull of adult o
(basal suture closed; teeth moderately worn) collected at Medje, Belgian
Congo, July 1, 1914, by Herbert Lang and James E Chapin. Orig. No.
2442.”

Crocidura congobelgica Hollister

Crocidura congobelgica HouuisTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV,
p. 670, Pl. vm, fig. 6 and PI. rx, figs. 3, 3a (skull).

“Type, No. 48512, Amer. Mus. Nat. Hist., skin and skull of adult
o (teeth little worn) collected at Lubila, near Bafwasende, Belgian
Congo, September 20, 1909, by Herbert Lang and James P. Chapin.
Orie No. 122.”’

There are only two specimens of this species in the collection, the
type from Lubila and a specimen from Medje.

Sylvisorex gemmeus irene Thomas
Sylvisorexr gemmeus trene HOLuIstER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV,
p. 671. Table of measurements of 12 specimens.

Specimens, 19: Medje, 18 (including 4 young in alcohol); Faradje, 1.

20 Bulletin American Museum of Natural History [Vol. XLVII

Sylvisorex oriundus Hollister
Plate II, Figure 2
Sylvisorex ortundus HOLuisTER, 1916, Bull. Amer. Mus. Nat. Hist., XX XV, p.
672, Pl. vu, fig. 7 and PI. 1x, figs. 4, 4a; Pl. x1, fig. 1 (animal).
“Type, No. 48554, Amer. Mus. Nat. Hist., skin and skull of adult 9
(teeth little worn) collected at Medje, Nava River, Belgian Congo, May
20, 1914, by Herbert Lang and James P. Chapin. Orig. No. 2368.”

Scutisorex congicus Thomas
Plate III

Scutisorex congicus HOLLISTER, 1916, Bull. Amer. Mus. Nat. Hist., XXXV, p.
673, Pl. x1, fig. 2 (animal). Table of measurements of 15 specimens.

Scutisorex congicus ALLEN, 1917, Bull. Amer. Mus. Nat. Hist., XX XVII, pp.
769-784, figs. 1-8, Pls. Lxxxrx-xcu. Skull and skeleton (Pl. xct, animal). Scuti-
sorex raised to the rank of a subfamily Scutisoricine.

Scutisorex (congicus) ScHULTE, 1917, Bull. Amer. Mus. Nat. Hist., XX XVII,
November 26, pp. 785-792. Thelumbar vertebre of Scutisorez.

Specimens, 44: Bafwabaka, 2 (skin and 1 complete skeleton);
Medje, 42, including 1 in alcohol and 5 more or less complete skeletons.

The highly specialized vertebral column of Scutzsorex has been made
the subject of two special papers already published in this Bulletin (loc.
cit., supra), one of them, by the author of the present paper, on the
remarkable specialization of the vertebral column—unique, or without a
known counterpart, in mammals—with numerous illustrations; the
other, by Dr. H. von W. Schulte, on the lumbar vertebre from the mor-
phogenetic viewpoint. In order to emphasize the taxonomic importance
of this surprising specialization the genus Scutisorex was raised to sub-
family rank under the name ScuTisoriciInm. To the first of these
papers Mr. Lang contributed several pages of field notes.

MACROSCELIDIDE

Rhynchocyon stuhlmanni stuhlmanni Matschie

Rhynchocyon stuhlmanni MatscuHin, 1893, Sitzber. Gesells. naturf. Freunde
Berlin, pp. 66-68. Andundi, Semliki River, two specimens, adult and young.

Rhynchocyon stuhlmanni nudicaudata LYDEKKER, 1906, Proc. Zo6l. Soc. London,
April 1907, p. 995. Mawambi district, Ituri Forest, Belgian Congo. One specimen.

Represented by 5 specimens: Penge, 2 (1 skin and skull; 1 alco-
holic), April 21, 1914; Avakubi, 2, November 13, 1913, and May 22,
1914. All are females, of which 2 are adult and 1 with the milk denti-
tion. Also a foetus in alcohol.

The Museum Collection contains also an unsexed specimen of this

form from the Budongo Forests, east of Lake Albert; February 1911.

1922] Allen, Congo Collection of Insectiwora 21

The collectors’ measurements of the two adults are: total length, 515
mm. (Penge), and 501 (Avakubi); head and body, 268, 259; tail, 247,
242; hind foot, 84, 84; ear, 30, 30. Skulls: total length, 69.5 (Penge),
—(Avakubi); condyloincisive length, 62.3, 65; zygomatic breadth, 36, 35.
They thus agree in measurements with average specimens of R. s. claudz
from localities farther west, as recorded below (Tables 1-4, pp. 23-26).

This fact has, however, little significance since the range in size of
adults of R.s. claudi covers all forms of the genus Rhynchocyon of which
measurements have been published. In coloration they closely resemble
extremely dark examples of claudz, from which they are not satisfactorily
distinguishable. Placed at the end of the dark series of claudz, they
completely merge with it. It seems preferable, however, to recognize
them as a darker geographical race of the same specific group.

Rhynchocyon stuhlmanni nudicaudata Lydekker, however, based on a
single specimen from the Mawambi district of the Ituri Forest, seems
scarcely entitled to serious consideration. The description indicates
that the type was not unlike the dark phase of the R. stuhlmanni group,
with which the author was at the time wholly unacquainted except
through the description of stuhlmanni. The “generally dark color and
wholly white tail’’ are not distinctive in view of the variations shown, and
described below, in the claudi series; nor are there any geographical
reasons that would seem to require its recognition, the type locality of
nudicaudata being less than fifty miles southeast frog Penge, in the same
environment as the latter, and represented in the present collection by
specimens of stuhlmanni, while the type of stuhlmannii came from a
locality equally near that of nudicaudata. The characters of naked ears
and tail, dwelt upon as important distinctions, have no real significance,
as such conditions are not infrequent in the claudi series; while a white
tail, at least in dry skins, is a prevailing condition. The hairs of the tail
are also so minute that they are often apparent only on very close in-
spection, giving the impression of a naked tail, especially in comparison
with examples of the czrnez group, with which the author compared his
specimen.

Rhynchocyon stuhlmanni claudi Thomas and Wroughton
Plate I; Text Figure 1

Rhynchocyon claudi THomas AND WrouGuHTON, 1908, Ann. Mag. Nat. Hist., (7)
XIX, May, p. 370. ‘‘Beritio, Welle River.”’

Rhynchocyon claudi Tuomas, 1915, Ann. Mag. Nat. Hist., (8) XVI, November,
p. 470. Medje, 1 specimen; Poko, 12 specimens. |

22 Bulletin American Museum of Natural History [Vol. XLVII

Represented by 105 specimens, of which 99 are skins with skulls; 6
foetal and young specimens in alcohol, and several skeletons, collected as
follows:

Medje, 20: May and August-October 1910; November 8, 1913;
March 17-20, 1914.

Nala, 1 (alcoholic): July 1913.
Akenge, 5: September 29-October 19, 1913.
Niapu, 79: November 8—December 26, 1913.

The 99 specimens represented by skins and skulls consist of 56 males
and 43 females, of which 76 are adult and 23 more or less immature. The
latter range in age from one specimen in which the milk teeth had not
pierced the gum to those with the deciduous dentition fully developed (a
series of 11 specimens), and the other 12 specimens fully illustrate the
transition from the deciduous to the permanent teeth. It has hence
seemed desirable to utilize this abundant material for the illustration of
the tooth development of this interesting genus of insectivores. (Text
figure 1, stages 1 to 8.) | |

The large series of adults from Niapu (43 males, 25 females) affords
the basis for a study of sex, age, and individual variation. In the fol-
lowing tables (Tables 1-3, pp. 23-25) the external measurements, carefully
taken by the collectors before skinning, have been combined with three
measurements of the skull (total length, condyloincisive length, and
zygomatic breadth). Table 1 gives the measurements of the males,
Table 2, of the females, and Table 3 is a summary of Tables 1 and 2.
In these tables the specimens are arranged in four categories, according
to age as indicated by the amount of wear shown by the teeth, the
purpose being to determine the influence of age upon the general size of
the animal after the permanent dentition has been fully acquired. Table
4 is designed to show the correlation of growth with the tooth develop-
ment. Of the 18 specimens included in this table, 5 are from Medje and
13 from Niapu, those from Medje being indicated by an asterisk.

Sexual Variation

There is no appreciable difference in size or coloration due to sex.
The average total length (tip of nose to end of tail) in 43 adult males is
515 mm., in 25 adult females, 516 mm. The average total length of the
skull for the same specimens is, males 68.1, females 68.5; condyloincisive
length, males 62.3, females 63.5; zygomatic breadth, males 36.3, females
36.2.

TasLe 1.—EXTERNAL AND CRANIAL MEASUREMENTS OF 43 ADULT MALEs oF Rhyn-
chocyon s. claudi, FROM N1APu, BELGIAN Conco

External Measurements Cranial Measurements
Head é Condylo- | Condi-
Cat. No. aire and Tail RE | Ear ae ee Pee tion of
Body Length Teeth
49442 500 265 235- |- 84 29 64.8 64.1 33.5 Unworn
49443 509 269 240 84° 31 67.0 61.6 36.2 ee
49444 505 265 240 86 ol 67.3 62.1 35.7 a
49445 | 556 291 265 89 | 34 67.3 63.5 36.3 x
49446 | 480 260 220 Sly | Si. hh 6526)) +6077 S5 ai fg
49447 482 250 232 83 32 67.2 60.9 36.4 2,
49448 531 286 245 86 | 33 66.1 61.4 34.8 a
49456 540 277 263 85 31 GS pi ObTo 36.5 oe
49458 900 303 202, 87 al == at 30.6 zs
49459 521 281 240 88 30 68.3 62.9 30.3 -
49461 504 206 Zaz, 90 31 Of 1 63.6 37.5 Ze
49473 6912 275 Zan 83 31 66.6 62.5 35.0 o
49474 458 260 Sas 83 3l 62.2 61.4 36.9
49475 469 244 225 82 31 66.9 60.8 34.3 i
49477 522 PA 249 88 31 66.7 60.7 34.4 oe
49478 510 278 232 87 jl 67.7 62.0 35.0 iS
49489 465 242 223 83 30 66.1 6E.2 33.6 ro
49492 496 259 237 86 31 67 .3 61.5 30.2 oN
49495 512 270 242 87 30 68.5 62.5 35.6 zs
49497 2 |) O74 248 86 30 68.5 62.9 36.8 +
49506 502 259 243 87 31 68.5 63.3 36.0 of
49509 518 274 244 84 30 66.8 61.6 aors -
49512 502 267 235 87 30 69.2 62.0 36.6 o
49515 514 275 239 85 31 68 .4 63.1 36.1 "
49527 | 516 262 254 86 29 66.3 G2E2 36.3 -
49449 Tw Dak 246 84 ol 69.0 61.9 36.4 | Shghtly
worn
49450 535 290 245 88 31 70.8 63.6 37.4 .
49462 527 275 252 90 31 68.3 62.4 36.9 en
49466 508 268 240 88 31 67.9 63.2 — re
49471 532 279 253 86 33 67.9 62.4 36.5 7
49496 5004 269 235 84 30 68.3 61.5 35.6 -
49451 5I2.-| 266 246 83 33 10.5 63.3 36.2 Much
worn
49452 535 287 248 89 31 66.7 63.6 38.0 es
49463 537 273 264 86 32 67.1 63.1 ote er
49482 510 267 243 84 32 68.4 62.1 36.3 Fa
49455 | 485 261 224 89 32 | —— | 63.0 38.1 | Greatly
worn
49470 520 291 229 87 32 67.7 63.1 37.9 .
49476 Syed 273 249 88 ol 69.2 62.7 37.2 i
49481 515 270 245 85 29 69.4 62.6 36.5 x
40491 505 270 235 83 29 67 .9 61.6 36.3 ef
49516 500 262 238 82 ‘30 65.6 60.1 36.5 *
49524 530 274 | 256 91 32 69.2 65.0 37.5 ~

24 Bulletin American Museum of Natural History [Vol. XLVII

TABLE 2.—EXTERNAL AND CRANIAL MEASUREMENTS OF 25 ApULT FEMALES OF
Rhynchocyon s. claudi From Niapvu, BELGIAN Conco

External Measurements Cranial Measurements
Head | : Occipito-| Condylo- Condi-
Cat. No. oe and : Tail ae Kar | ee | ee Ra i
Body | Length | Length Teeth
49460 | 532 281 251 85 3l 68.4 63.6 36.7 |Unworn
49464 | 530 272 258 86 jl 67 .6 62.3 34.5 i
49479 | 529 271 258 85 32 67.6 61.8 34.3 -
49480 | 516 279 | 237 88 dl 69.5 | 63.0 36.0 %
49485 | 525 266 259 84 30 67.9 | "6323 )aeeet ‘
49486 | 522 276 246 89 | 29 66.9 | 62.1 — :f
49487 | 523 274 249 87 32 68 .0 63.5 36.5 “
49488 | 514 276 238 85 30 68.4 | 63.7 30.4 “
49490 | 535 283 252 84 29 66.1. | 62.2 36.6
49500 | 522 277 245 85 30 68.8) =| 76226 36.1 -
49504 | 517 262 259 85 dl 68.8 | 63.6 34.2 S
49507 | 5dl1l 270 241 | 86 dl 6755) | Oh 33.6 Reset
49511 530 217 253 87 30 70.6 | 6353) heaoma bs
49526 | 513 Die, 241 86 3h. | 68ai he Oama 35.6 ee
49502 | 512 201 241 86 dl 67.6 62.5 35.8 (Slightly
| worn
49508 | 492 255 237 81 ol 66.4 61.3 | aor .
49453 | 517 277 =| ~=240 85 dl 68.5 63.1 | 36.5 | Much
| | | worn
49469 | 528 29a NWN e2ao 87 dl 70.6 64.0 36.7 ne
49483 | 539 294 | 245 86 dl 69.6 , 63.6 | S583 ci
49494 | 511 266 | 245 87 dl 69.2) 63.1 owt vi
49501 | 540.) 292 | 248 |. 87 | 3L | 68.0.) 635 “| aasne
49503 | 520 278 | 242 85 32 71.1 | 64.8 37.4 |Greatly
| | | | | worn
40454) | 5327") 285 | 247 | 86 | Sl 69.00 63! aoe aaeoes :
49472 | 505 272 | 233 83 32 68.8 | G61.8= esuen i
49525 | A99: 9 273) 12226 | 88 33 69.1 64.0 36.2 re

1922] Allen, Congo Collection of Insectivora 25

TABLE 3.—SUMMARY OF MEASUREMENTS OF 43 MALES AND 25 FEMALES GIVEN IN
TABLES 1 AND 2

Unworn|Avg. | 0°25 | 508.0 | 269.7 | 240.5 | 85.4 | 30.8 | 67.2 6
SeeeiMane | ot25 |.458 | 242 (| 220 81 29 64.8 | 60.7 | 33.5
5

SeeeiMiax th Gt25 | 556 | 303. | 265 || 90.) 84 | 69.2) | 64.1) 37.
Slightly | |
worn |Avg. | 3 6 | 519.3 | 274.0 | 245.3 | 86.1 | 31.6 | 68.9 | 62.5 | 36.5
Seine ct 6 | 504 | 266° | 235) |'83 (| 80) | 67.8 61.5 | 35.6
Bee Mee oN 15385 «| 200.) 253 | 90. | 38u 170-8: |, 6386 13704
Much | | | | |
worn |Avg. | o 4 | 523.5 | 273.2 | 250.2 | 85.5 | 32.0 | 68.2 | 63.0 | 36.9
pee tie) c? 4) 510 266 | 243 1°83 =| 31 1 66-7 | 62.16 136.2
Pee Mex | ot 4 | 535. | 287 | 264 89 33 70.5 | 63.6 | 38.0
Greatly | | |
worn |Avg. | 7 | 511.0 | 271.6 | 239.4 | 86.3 | 30.7 | 68.2 | 62.6 | 37.1
Seen st 7 | 430 | 261 | 224 | 82° | 29 165.6 | 60:1 | 36:3
Seiler) ot 7 | 485. | 201. | 256 «|.91 -| 32 1-69.42) 65.0.1 38.1
Unworn|Avg. | 914 | 522.8 | 278.6 | 248.6 | 85.9 | 30.6 | 68.1 | 64.2 | 35.5
ein: | O14 | 511 22 e28n ty | S4e 29 66.1) s6128) | San
pee Mase O14) 535 || 283 -| 259, | 89 + | 82° 170.6 63.7 | 36.7

Slightly | | | | | | |
worm jAve, | © 2) 502.0) 268.0 | 239.0 |-88.5 | 31.0 | 67.0 | 61_9 | 35.6
Se Min. | -Ox2 | 492 255 1) 223% 81 66.4 | 61.3 | 35.8
fe Max. | 0 2 | 512 2h DATS 86 | Si 67.6 | 62.5 | 35.4
Much | | | | |

iss)
—_

worn |Avg. | 9 5 | 525.8 | 281.7 | 242.5 | 86.2 | 31.2} 69.5 | 63.5 | 36.7
ee Mit O75 | ott 266° 235185 | | Sh | 6820 6881 P3573
em iar 540) (204 | 248 | 87 1 32) oA 64-00 8704
Greatly | | | | | | |
worn |Avg. | 9 4 | 512.0 | 276.7 | 235.3 | 84.0 | 32.0 | 69.0 | 63.4 | 36.2
SeeiMan | o 46) 409 “279° | 996 «| 8B | 81 168.8 | 61.8 | 35.7
PemaMax. |) G.4.| 532) ) (285. |\247 ~ | 86 “| 38>] 69.1 | 64.8 | 36-8

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1922] . Allen, Congo Collection of Insectivora 26

Age Variation

CoLORATION.—Coloration is only slightly affected by age. In
young specimens in the first pelage, the tones are practically the same as
in adults; the light and dark markings of the upperparts are not quite so
sharply defined, but the pattern is strictly the same. In very old speci-
mens the dorsal pelage has sometimes a more grayish cast than is usual
in younger animals, due perhaps to less prompt renewal. As will be
shown later, the wide variation in color seen in a series of specimens has
no relation to sex or age.

Si1zE.—Tables 1-4 have been compiled with special reference to the
effect of age upon the general size of the animal and upon the size of the
skull, since species and subspecies are sometimes based on adolescent
specimens, and frequently on ‘‘young adults.”” Table 4 shows that in 6
specimens having only the full deciduous set of teeth the total length
(tip of nose to end of tail vertebrz) ranges from about 440 to 460 mm.
as compared with the average adult length of about 515 mm., and a con-
dyloincisive length! of about 50 to 60 mm., as compared with about 63 mm.
in middle-aged specimens. During the period of the replacement of the
deciduous by the permanent teeth the size increases to about the mini-
mum for adults, the total length averaging about 500 mm. and the
condyloincisive length about 60 mm.

‘The average total length of the animal in 38 specimens (24 males, 14
females) with unworn teeth is 514 mm. (males 509, females 524); aver-
age condyloincisive length of skull, 62.3 (males 62.1, females 62.8).

In 8 specimens (6 males, 2 females) slightly more advanced in age
(the teeth appreciably worn) the average total length is 516 mm. (males
520, females 501); condyloincisive length of skull, 62.3 (males 62.5,
females 61.5). In this case the number of specimens is too small to be
satisfactory, especially in relation to sex difference in size.

In 9 specimens (4 males, 5 females) still older (teeth much worn), the
average total length is 524.4 mm. (males 523, females 527); condyloin-
cisive length of skull, 63.27 (males 63.25, females 63.5). Again the
series is too small for satisfactory results, but is not wholly without
interest.

The old-age (senile) series 1s represented by 11 specimens (7 males,
4 females). The average total length is 512 mm. (males 511, females 514);
ee length, 62.9 (males 62.6, females 63.5).

1The condyloincisive length is a better standard than total length of skull, the ossificaticn of the
nasal cartilage being a variable element, sometimes terminating at cr a little tehind the tip of the
premaxille but usually extending several millimeters keyond this point. Hence, tctal lergth erd oc-
cipitonasal length are undesirable measurements for skulls of Rhynchocyon.

28 Bulletin American Museum of Natural History [Vol. XLVITI

The results of the foregoing analysis of variation in size as affected
by age and sex are collated in the following tabular résumé (Table 5).

The results derivable from the above tabulation would possess
greater interest if the number of specimens in each of the six categories
had comprised a more nearly equal number of specimens—if each had
been as large as in Table 3. It seems safe to assume (1) that size is not
diagnostic of sex, although the above statistics indicate a slight superiority

TABLE 5.—RELATION OF AGE AND SEX TO SIZE

C Ley | No. of tome aoe
‘ondition of Teeth snc cnuees Length aie.
| Animal Skull
1. Entire milk set only | 6 451 56.3
2. Entire milk set plus me more or less developed 5 | 501 60.5
(38 514 62.3
3. Permanent set, unworn {2497 |. 209 62.1
149 524 | 62.8
8 516 62.3
4, Permanent set, slightly worn 6c 520 62.5
29 501 615 =
9 | 524.4 63.27 .
5. Permanent set, much worn . 4 | 823 Goes
il 5 527 63.5
11 512 jc 62-9
6. Permanent set, greatly worn TS Le ial 62.6
Hen ene yb ell! 63.5

for the females. The largest specimen of the entire series is a ‘‘ young
adult”? male (No. 49445), with a total length of 556 mm., a tail length of
265, length of hind foot 89, and condyloincisive length 63.5, dimensions
not reached by any female, except the skull length in a few old females,
which again is exceeded by a few old males. (2) There is a slight increase
in size, both externally and of the skull, in the old-age period, but insuffi-
cient to antagonize the selection of young adult specimens as types of new
forms, since individualism in any age class more than bridges the differ-
ences that can properly be ascribed to age after approximate maturity
is reached.

1922] Allen, Congo Collection of Insectivora 29

Individual Variation:

SizpE.—As already noted incidentally above, the variation in total
length (tip of nose to end of tail) in the series of 25 young males with un-
worn teeth from Niapu covers the entire range of variation in the whole
series of the 68 adults from Niapu, all of which were taken within a period
of about six weeks in November and December of the same year, and all
within a radius of about six miles in strictly uniform environment.
Leaving out of consideration a single specimen (No. 49474, o”), obviously
a dwarf, the average total length is 510 mm., the two extremes being 465
and 556, a difference of 91 mm., 17 per cent of the mean. Even this is
exceeded in the old-age series of 7 males, where the range is 19.6 per
cent. This illustration apples equally to length of tail, where the range
of variation is 18.7 per cent of the mean, but not to hind foot and ear,
where the range is respectively 10 and 2 per cent. It is also much less in
the skull, in which the mean condyloincisive length in the 25 young
adults in question is 62.1 mm., and the extremes 60.7 and 64.1, and the
difference 3.4 mm., or only about one-half of 1 per cent. This, however,
is nearly equal to the variation due to age, where the average condylo-
incisive length in the old-age series of 11 specimens is 62.9 mm. (minimum
60, maximum 65 mm.). The variation in zygomatic breadth parallels
that of the skull length.

CoLoraTion.—Rhynchocyon s. claudi may be said to have, ina general
way, a light phase and a dark phase of coloration, but a large proportion
of the specimens in the present large series are in such varying degrees
intermediate that no line of demarcation can be even approximately
assigned. As the extremes of light and dark specimens belong to the
same sex and prove to have been taken on the same day at the same place,
it must be assumed that this wide range of color variation is purely in-
dividual. Yet, should single specimens of the extremes of the light and
dark types of coloration be received by a systematist from even the same
locality, he might be pardoned for considering them as nameable forms.
Some of the East African forms of Rhynchocyon have been found to be
notably prone to melanism, but among the hundred examples of the
claudi type collected by the American Museum Congo Expedition not
one shows such tendency, notwithstanding the large amount of color
variation they present.

The light or reddish phase (Plate I, upper figure) may be indi-
cated as follows, beginning with the ventral area:

Chin, throat, fore neck and pectoral region entirely and nearly uniform buff,
varying from pale buff to ochraceous buff (in different specimens), abruptly con-

30 Bulletin American Mi seum of Natural History [Vol. XLVII

stricted at axille and pectoral area to about the median third or fourth of thorax,
thence expanding to cover the lower abdomen and inside of thighs, usually darkest
on middle of breast and lighter on throat, middle of thorax and mid-lower abdomen.
In extreme specimens this portion of the ventral surface has a decided rufous tone.
Sides of head from base of rostrum, expanding upward to enclose the ears, sides of
neck and sides of body to base of tail (encroaching deeply on sides of abdomen and
nearly meeting over thorax), brownish rufous or chestnut slightly varied with black-
tipped hairs. Top of head and mid-region of back to base of tail more varied with
black-tipped hairs, which from the withers posteriorly take the form of four longi-
tudinal blackish bands, which from middle of back to base of tail are broken by four
or six transverse rows of whitish spots, which vary in tone (in different specimens)
from clear white to pale buffy white. Over this area the general effect is that of
alternating transverse rows of rather sharply defined black and white spots, about
five of each being rather distinctly defined, with an additional posterior row of two
white spots at the base of the tail, and an ill-defined anterior row of small, less dis-
tinct, whitish spots. There is also a tendency to an additional lateral row of indis-
tinct or subobsolete whitish or pale buffy spots on each side of the usual four distinct
median rows of spots. Counting all the rows of white or whitish spots they form
six longitudinal rows, the outer rows separated from the others by dark chestnut
instead of blackish intervals.

No. 49463, Niapu, November 24, 1913, adult o, and No. 49477, Niapu,
December 1, 1913, adult o, may be taken as typical of the light or reddish phase.
In general tone No. 49477 is lighter, with the dorsal spots clearer white, than No.
49463.

The dark phase (Plate I, lower figure) may be thus indicated:

Light portion of the underparts much paler, or faintly yellowish white; the sides
of head, neck, and body dull dark brownish, almost without trace of rufous except
around ears and on sides of neck; top of head and mid-region of back grizzled yellow-
ish gray with most of the hairs broadly black-tipped; the back from the posterior
part of thoracic region to base of tail with deep black predominating, the whitish
spots reduced in size and usually rather clear white, and the longitudinal and trans-
verse bands indistinct or blended into a black or blackish general ground color,
the black most concentrated along the median line.

This phase is typically represented by No. 49487, Niapu, December 4, 1913, adult

Q. No. 49490, adult 2, same locality and date, has more rufous suffusion on sides of
neck, nape, and shoulders.

Each phase is typically represented by both males and females taken
on consecutive days, or sometimes on the same day, at the same locality.
Other specimens collected actually or approximately at the same date
and place, equally representative of both sexes and strictly comparable
as to age, fill every gradation between the two extremes. Hence the
types of coloration above described can scarcely be considered as repre-
senting respectively a definable red and dark phase, but merely the ex-
tremes of a wide range of purely individual variation, shown in the
accompanying colored plate.

1922] Allen, Congo Collection of Insectivora 31

CRANIAL VaRIATIONS.—Matschie! and others apparently believe |
that the relative length of the frontal and nasal sutures is of specific value
in Rhynchocyon. Specimens of Rhynchocyon s. claudi in our series show that
the frontal suture may be as long as, or longer or shorter than, the nasal
suture, and in some cases one of the nasals is fully 4 mm. shorter than
the other. ‘The following measurements? illustrate variations in speci-
mens taken in the same locality at Niapu:

Frontal Suture Nasal Suture

No. 49445 of 24 mm. 24 mm.
49479 9° DAT PAS) Tone oe 19 mm.
49443 o 27 mm. 20 mm.
49459 of 28 mm. 20.5 mm.
49448 of 7s eNO 25 mm.

Rhynchocyon claudi Thomas and Wroughton was based originally
on a specimen in the light or reddish phase of coloration from Beritio,
near Angu, on the Uele River. Later a single specimen from Medje and
twelve others from Poko were referred to this species by Thomas.
The present collection contains 20 specimens taken at Medje, 5 col-
lected at Akenge, and 79 at Niapu. The two last-named localities
are within about thirty miles of Medje and Poko and have the same
environment. There can be little doubt therefore of the correct refer-
ence of all these specimens to R. s. claudt.

DENTITION OF Rhynchocyon
Text Figure 1

The present large series of skulls of Rhynchocyon's. claudi affords
material fully disclosing the character of the dentition of Rhynchocyon
from its early stages to old age. In the youngest skull (No. 49434—see
Table 4, p. 26) of the series the teeth are wholly enclosed in the gum;
in a slightly older specimen (No. 49427) the tips of the principal cusps
of the deciduous teeth (canines and premolars 2, 3, 4) have broken
through. Other specimens, more advanced, show the gradual develop-
ment of the milk teeth and the order of their displacement by permanent
teeth. In a succeeding table (Table 5, p. 28) measurements are given
to show the correlation of the size of the individual with tooth develop-
ment, from the stage just prior to the appearance of any of the teeth

11893, Sitzber. Gesell. naturf. Freunde Berlin, p. 6
2Other cranial variations are indicated in the ae of measurementsand need not be especially
emphasized, as they present no unusual features.

OZ Bulletin American Museum of Natural History [Vol. XLVII

above the gum to full maturity of the permanent set. Several stages of
development are also shown in the accompanying text figure (Stages
1-8).

Deciduous Dentition

The milk dentition, strictly construed, consists of 24 teeth: I53,
CH, P$3=1)=24. The first premolar (p ©) is not present till later
and has no successor.

Upper oR Maxititary Sreries.—The single upper incisor (i? by
position) is a minute spicule inserted at the extreme posterior border of
the premaxilla and has no successor. Although small and frail, it often
persists through life, being frequently present in the senile stage. In 46
adult skulls, taken at random for the investigation of this point, 15 (33
per cent) were found to retain one or both upper incisors, both being
present in 6 skulls and one in each of 9 skulls, most frequently on the
right side. When these teeth are absent their alveoli often remain,
indicating the recent presence of the teeth.

The canine is a small bicuspid tooth, with a slender-pointed central
cusp, and a small slender-pointed posterior cusp, about one-third as
high as the main cusp. The canine is shed at the same time as the pre-
molars, but its successor is long in maturing, and, when fully developed,
is long, slender, and saber-like. The second, third, and fourth premolars
(dp?, dp’, dp*) arise simultaneously. Dp? has a basal length slightly
exceeding its height, with two pointed cusps, the anterior one consider-
ably exceeding the other in size and height, and a low anterior and a low
posterior cusplet, both arising from the cingulum. Dp? is subtriangular
in basal outline, the anterior half narrow, the posterior broad, with a
main central cusp, a smaller one behind it, and a still smaller one in
front, on the cingulum. There is also a low, broad postero-internal cusp,
and, behind this, a slight cusplet from the cingulum. Of these five cusps,
three are external and two internal. Dp?’ is subquadrate and distinctly
molariform, with four prominent cusps, the outer much higher than the
inner, the four cusps collectively enclosing two deep basin-shaped
cavities. There is also an anterior cusplet from the cingulum.

Lower OR MANDIBULAR SERIES.—The anterior four milk teeth in
the lower jaw are all incisiform, similar in size and general form, and have
their axes directed forward. The two middle teeth are tricuspid, the
first and fourth bicuspid. They are separated from dp: by a long convex
diastema. The first three incisiform teeth are shed singly at intervals.
The three posterior milk premolars (dps, dp3, dps) increase successively

1922] Allen, Congo Collection of Insectivora 393

in size, dp2 being less than half the size of dps, and dps is less than one-
third the bulk of dps. Dp2 has a high-pointed central cusp and a small,,
low, sharp-pointed one before and behind it, and a cusplet on the posterior
cingulum. Dp; is similar in structure to dps, but is a much larger tooth.
Dp. consists of two sections, each of which encloses a deep basin from the
borders of which arise four cusps, of which two are antero-external,
the other two internal, one of which is median and the other posterior.
The medio-internal cusp is usually minutely bipointed when unworn.
The above conditions are represented, essentially or exactly, by 7
skulls (Nos. 495238, 49486, 495138, 49514, 49510, 49499, 49493, of Table 4).

Permanent Dentition

The permanent dentition comprises 36 teeth: I33, C7, Pi, M
= =5=36. In this enumeration the minute upper incisor is assigned
as a permanent tooth, although, as already explained (p. 32), it is often
absent in adults, though frequently persisting through life, and has no
successor. The first premolar in both jaws has also no successor and is
developed later than the other premolars which have successors. _ |

Uprrr oR Maxintuary Sertes.—The canine is a long, slender,
laterally compressed, 2-rooted tooth, with a conspicuous longitudinal
groove on its antero-internal face. The first premolar (p') does not
pierce the gum till the milk premolars (dp?, dp’, dp*) are fully developed
and functional, and has, as already said, no predecessor. It is a small
unicuspid, 2-rooted tooth, about as long antero-posteriorly as high. It
is separated from both the permanent canine and the permanent p? by
diastemata nearly equal in length to the basal length of the tooth. P?
and p* are similar in form to their respective predecessors, from which
they differ mainly in larger size. P*is more completely molariform than
dp‘, and differs from m! only in being larger and slightly more quadrate.
M! is subquadrate, the anterior half of the tooth broader than the pos-
terior half, with higher cusps, which are situated at the four corners of
the tooth. M? is trigonal, with three cusps, and is about one-third the
size of m*. Thus, in the permanent dentition, the last three maxillary
teeth are typically molariform and, on the basis of structure and posi-
tion, would be classified as molars, but the first one of the three has a
milk predecessor.

Lower oR ManpiBuLAR SuRrES.—The three permanent incisors
all have bifid crowns, are close-set, directed obliquely forward, and differ
from their predecessors mainly in their larger size. The canine is a small
2-rooted tooth, separated from i3 and p, by slight diastemata. Its axis

A.M.49513

A.M.49498

Fig. 1. Development of the dentition in Rhynchocyon stuhlmanni claudi
Thomas. All X%; a and 5, side views; c and d, crown views. From specimens taken
in the Belgian Congo by The American Museum of Natural History Congo Expe-
dition, 1909-1915.

Stagel. Milk teeth breaking through gums. No. 49518, 9 juv., Medje, June 3, 1914.

‘ ga 2 Milk teeth more advanced; alveolus of p! fissured. No. 49437, 2 juv., Medje, Septem-
er 8, K

Stage 3. Milk teeth fully developed, and p! nearly so; convex diastema where pi is forming be-
neath. No. 49513, ojuv., Niapu, December 17, 1913.

Stage 4. Upper milk dentition p!, and crown of m! just through gum; lower jaw: milk teeth, p1
Ny oracvanced, mi half up, and tip of permanent i: in sight. No. 49498, @ juv., Niapu, December 6,
1913.

ss r

A.M.49503

(Fig. 1 continued)

Stage 5. First molarfully up and second molar half up in both jaws; permanent premolars visible
between the roots of their predecessors; permanent lower incisors and lower canine fully developed.
No. 49484, 2 juv., Niapu, December 4, 19138.

Stage 6. Slightly more mature, the molars being full-grown, but premolars of permanent set still
covered by the milk teeth in both jaws; caniniform pi nearly mature; upper milk canine has been
shed. No. 49468, o&, Niapu, November 25, 1913.

Stage 7. Mature permanent dentition, but upper canine not fully grown. No. 49486, 2, Niapu,
December 6, 1913.

Stage 8. Senile stage toshow wear of teeth. In old age the first premolar becomes greatly worn in
both jaws, sometimes only the roots of p! remaining. No. 49503, old 9, Niapu, December 8, 1913.

36 Bulletin American Museum of Natural History [Vol. XLVII

is directed forward, as is the case with the incisors, which it exceeds but
little in size, its blunt-convex crown not rising above the crown surface
of the incisors. P; is a 2-rooted unicuspid, perfectly caniniform in struc-
ture and function; its height is about twice its basal length, or about
twice the height of pz and ps, and it is a persistent tooth of the first set.
Ps, ps, and p, are respectively similar in structure to their predecessors
but more massive. P., conforms in plan with the molars except in having
an additional cusp on the interior border, making five cusps instead of
four as in the molars. It is thus a slightly longer tooth than m,. The ©
median internal cusp, when the tooth is unworn, is usually minutely
bipointed, as in its predecessor. My, and ms, are similar in structure, but
ms is only about half the size of M4.

Development and Replacement of the Deciduous Teeth

The condition of the teeth, as seen in the cleaned skull at the stage
when the crowns of the last three premolars in each jaw (ees aa ,
are fully excluded and the teeth have become functionally effective (skull
No. 49513), is as follows. The incisors and canines (both above and
below) are, like the last three premolars, fully developed and functional;

the extreme tip of p! is barely above the alveolar border and would be,

in most cases at least, still covered by the gum; dpe is not visible but

there is a narrow slit at the outer base of the convex diastema between
the canine and dpe; there is also a narrow opening in the alveoli of the
future molars, in each jaw.

Upper Trpetu.—Later (Fig. 1, Stage 4), after the skull has nearly
doubled its size, p! attains its full development. P! persists without
change except by wear, and, later in life, is the first tooth to become in-
efficient through excessive attrition. After considerable further increase
in the size of the skull the crown of the first molar is excluded and, as
soon as it has reached functional maturity, is followed by the second
molar. Not, however, till m? has become fully functional is there any
further change, when the crowns! of the permanent canine and the
permanent premolars (p?, p’, p*) can be seen between the roots of their
predecessors, but it is considerably later before the milk premolars are
finally shed and their successors are fully developed.

Lower Tretu.—tThe first change in the milk teeth of the mandible
occurs coincidently with the breaking of the crown of m, through its
alveolus and before the crown surface of the tooth is much above the

1In skull No. 49428 the crown of the right canine is double, consisting of two equal slender stiles in
close contact. e

1922] Allen, Congo Collection of Insectivora 37

alveolar border. At this stage (represented by No. 49499) the four incisi-
form teeth are still unchanged, but the tip of the caniniform p, is just
above the alveolar plane, or practically at the same stage as m;. At the
next stage (represented by No. 49498, Fig. 1, Stage 4), m; is about two-
thirds grown but not as yet at full height, and p;is evenly keeping pace with
it in development. None of the other milk teeth shows any indication of
approaching replacement. In skull No. 49428 this molar is fully de-
veloped, as is also p;, but the milk premolars (ps, ps, ps) are still firmly in
place, with no trace of their successors between their roots. The first
incisor (i;), however, has been renewed, and the crowns of the successors
of i2and i3can be seen pushing up at their inner bases, and the same
condition is also true of the canine. It would seem, therefore, that the
renewal of the milk incisiform series just precedes that of the milk pre-
molars. In No. 49484 a somewhat later stage is shown, in which m, is
not only fully mature but the crown of mz has reached about one-half its
fullheight. In this skull (Fig. 1, Stage 5) the permanent canine has attain-
ed to about half the height of its predecessor, coming up at its outside base,
and the crowns of the milk premolars can be seen between their roots,
these four teeth, like the same teeth in the maxillary series, developing
coincidently. The permanent incisors and the permanent canine are also
now fully matured.

To complete the series of illustrations a figure of the unworn denti-
tion of a youngish adult (Fig. 1, Stage 7) and of an old-age adult (Fig.
1, Stage 8) are here included. .

Nasilio fuscipes (Thomas)

Macroscelides fuscipes THOMAS, 1894, Ann. Mag. Nat. Hist., (6) XIII, January,
p. 68. “N’doruma [Doruma], Niam-Niam country (about 5° N. and 27° 30’ E.).”’
Based on a young female ‘‘ having still its milk-dentition in place.”

Represented by 30 specimens (28 skins and skulls, 2 in alcohol), col-
lected as follows: |

Niangara, 8: November 18—December 19, 1910.

Faradje, 21: February 8, 1910; February 20-May 6 (mostly Feb-
ruary 2—March 1), 1911; December 25, 1912; January 3-8, 1913.

Garamba, 1 (in alcohol): March 1912.

Males and females are equally represented: All are adult except 9,
, which are one-half to two-thirds grown, with the permanent dentition
incomplete. These form a series showing all the stages of change from the
deciduous to the permanent dentition, confirming Thomas’s determina-
tion of the dental formula in Macroscelides.'

11890, Proc. Zo6]. Soc. London, pp. 445, 446. Milk dentition of Petrodromus figured, p. 445.

38 Bulletin American Museum of Natural History [Vol. XLVII

The adult males have a short-haired, glandular pectoral area, the
short hairs of which, and the longer enclosing pelage, are cream-color,
due possibly to staining. In some males this area has an axial extent of
20 mm. or more and a breadth of 10 mm., thus forming a conspicuous
feature of the ventral surface, but it is less developed in females.

The Niangara series, taken at the close of the rainy season, are
rather brighter colored—more rufescent and less gray above—than those
from Faradje collected some two or three months later. The difference
is not wholly constant and may be seasonal, as some specimens of the
Niangara series are indistinguishable in coloration from some of the
Faradje specimens.

Collectors’ measurements of 7 adult males and 10 adult females from
Faradje:

Total Length Head and Body Tail Vertebree Hind Foot Ear
o& 204 (200-207) 115 (109-128) 91.3 (84-96) 29.3 (28-31) 21.0 (20-22)
® 210 (201-224) 120 (114-129) 91.0 (85-99) 29.2 (28-30) 22.2 (20-23)

Skulls, 5 males, 9 females, Faradje series:

Total Length Zygomatic Breadth
ao 34.4 (84 -34.7) IR OIG} 11)
Q 34.9 (386.6-36.4) 17.4 (16.6-18.1)

The minima are all from rather young adults, the maxima from
obviously very old skulls. The females show a slightly larger average
size than the males, both in external and cranial dimension.

These specimens agree satisfactorily with the description of the type
of fuscepes when those corresponding with it in age (‘‘a somewhat imma-
ture female. . . having still its milk dentition in place’’) are con-
sidered. Geographical conditions may be taken as confirmatory of this
identification, the type locality (Doruma) of fuscipes being about 100
miles north of Niangara and some 160 miles west of Faradje and
Garamba. These four localities, Mr. Lang assures me, are in regions of
similar environment. The present series, if here correctly referred,
shows that Macroscelides fuscipes Thomas belongs to the genus Naszlio
Thomas and Schwann! (type Macroscelides brachyrhynchus A. Smith),
the molars being 3. The type of fuscipes was so young that it could give
no hint of the number of the molars, but it is described as ‘‘most nearly
allied to M. brachyurus Boc.,” of which the author says: ‘‘Sa machoire
inférieure porte chaque cdté une molaire de plus, onze dents au leu
de dix.’”?

11906, Proc. Zodl. Soc. London, II, p. 578. .
2Bocage, 1889, Journ. Sci. math. phys. nat. Acad. Madrid, (2) I, No. 1, p. 24, Marco.

PLATE I

Rhynchocyon stuhlmanni claudi Thomas and Wroughton. ‘Two males, taken in
the same locality within a week, showing an extreme light rufous and an extreme dark
phase. Drawn by Charles R. Knight from skins, Amer. Mus. Nos. 49495 and 49477,
Niapu, December 1 and 6, 1913.

T F4V1g ‘ITATX “IO ‘HON ‘WW ‘V Niiaring

Prarn Tt
Fig.1. Crocidura jacksoni denti Dollman. Female adult, Amer. Mus. No. 48520,
Medje, May 28, 1914.
Fig. 2. Sylvisorex oriundus Hollister. Type. Female adult, Amer. Mus. No.
48554, Medje, May 20, 1914. Photographs from specimens in the flesh. Both
natural size.

Buuietin, A. M. N. H.

Vou. XLVII, PuatE

Puatre III

Scutisorex congicus Thomas. Male adult, Amer. Mus. No. 48475, Medje, May
30, 1914. Photograph from specimen in the flesh. Natural size.

ih nyo send taMe cate.

eS " i ss a ee ii pated Ac aS il aN aia.

WoOYV SNIL@T1og

III 224V1q ‘TIATX “TOA ‘HN

Puate- lV

Crocidura nyanse kivu Osgood. Male adult, Amer. Mus. No. 48501, Medje,
June 13, 1914. Photograph from specimen in the flesh. Natural size.

AI ®4V1Tq “TIATXi TOA ‘HON CIN ‘Y ‘Niwai1og

pt
iE
’

Sue va

| Ce SP ee SR ate te E Eexeiotoge | ee
On Oe Seite mentalis and its hes By: W.°E. Bivde Todd, 1916, 8

Bulletin, XXXV, ‘Art. 29, pp. 533-558, 4 text figures.

"The Distribution of Bird Life in ‘Colombia; a Contribution to a ‘Biological atk S

- Survey: of South America. ° By Grank M. Chapman, 1917, Bulletin, :

XXXVI, pp. 1-729, Pls. tx, 21 text figures.

* The: Classification of the Weaver-Birds. By J: ames~ Pp. Chapin 1917,

‘Bulletin, XXXVU, Art. 9, pp. 243-280, Pls. vi-x, 10 text figures.

_ Descriptions of New Birds from Santo Domingo and Remarks on Others in.

the Brewster-Sanford Collection. . DY: Frank M. Pectin ee 1917, Bulle-
tin, XXXVI, Art. 12, pp. 327-334. =

Field Notes on Molothrus bonariensis and M. badius. “By Leo. BE Miller :

++ 1917, Bulletin, XXXVII, Art. 22, pp. 579-592.

2 Besa ions of Four Birds from the Belgian Congo. By James P; Chapin, ae

2 A New Albatross from the West Coast of South hens By Robert. Gash:

me man Murphy, 1917, Bulletin, XXXVIL a 35, pp- 861-864, 1 text
figure.

ey ‘Study of the eGtiG Deca “By Robert Cacia ares 1918, De

~~ Bulletin; XX XVIII, Art. 4, pp. 117-146, Pls. 1-111, 7 text figures.

~The Geographical Distribution of Color and of Other Variable eects is

in the Genus Junco: a New Aspect of Specific and Subspecific Values.

~ By Jonathan. Dwight, M.D., 1918, Bullpen, XXXVI, Art. 9, PP. 269- .

- 300; Pls. xi—xmu, 5 maps.

" Destriptions of Proposed New Birds from. Perth Relive peeing. and
~ -. Chile,” By Frank M. Chapman, _ 1919, Bulletin, a Art, 5, Pee

323-333. °

_ Descriptions of Apparently New Birds ioc Bola Brazil, and Venezuela.

By Frank M. Chapman, 1921, Novitates, No. 2, pp. 1-8.

1921, Novitates, No.7, pp. 1-9.

ik Revision of Ailapetes gutturalis with Descriptions of Three N ew ioe By

Jonathan Dwight and Ludlow Griscom, 1921; Novitates, No. 16, pp. ae

_ Notes on a New Ox-pecker and Other. Little-known Birds of the Congo. By
- James. P. Chapin, 1921, ‘Novitates, No. 17, pp. 1-16.

= Descriptions of Proposed New Birds Ae Colombia, Ecuador, Peri, and

Brazil: By Frank M, Chapman, 1921, Novitates, No.18, pp. 1-12.
“Descriptions of Proposed New Birds oe Central America; with. Notes on

~ Other Little-known Forms. By Waldron DeWitt Miller and Ludlow fs

- Griscom; 1921, Novitates, No. Ob, pp. 1-13.

ee e ; A Review of the Divied Petrels, Contributions fram the Brcwsicie cantor’
- Collection. By Robert. Cushman Murphy and Francis Harper, 1921,

Bulletin, XLIV, Art. 17, pp. 495-554, Ps: XX-XXIV, 7 text figures. ~

-Deseriptions of Proposed New Birds from Brazil, Paraguay, and Argentina. -
By George K. Cherrie and (Mrys. .) Elsie M. B: Reichenberger, Bee

Novitates, No. 27, pp. 1-6.:

- Expediti ion of 1916-1917; By Outram Panes; 1921, onienrne XLIV,
_ Art. XX, pp. 575-612. :

The Distribution of the: Swallows of the Gti Pyannketitin = By Prank ;

M. Chapman, 1922, Novitates, No. 30; pp. 1-15.

~ Deseriptions of Apparently New Birds from Colombia, _Ecuador, dnd
~ Argentina. By Frank M, Chapman, 1922, Novitates, No, 31, pp. 1-8. 7

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= Volumes xXx. The = Sree Lik — record of the progress
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CA OE OS Fy § OR ee) ek

os Se ee ee ine Lom, Epon |
O oe The following a are . the: yore recent ‘papers « on M AMMALOGY AN
oe (Ont cone Orders should be addressed, Library, ‘The Ame
Museum of Natural tatty 4 7th St. and ess ark
ee Oty. ae cae ee 2
pie a ae - Maonstocy 2
_ . om Note o on ‘she Santer euch: of Scutisorex Thomas. _ By rE
ee =e - Schulte, 1917, Bulletin, XKXVU, Art. 29, pp. 785-792. e
8 os aS External Characters, - Skelet al “Muscles, and Peripheral Nerves:
oe a _Kogia breviceps (Blainville). “By H. von 'W. Schulte and M. de F Forest.
eS ~ Smith, 1918, Bulletin, XXXVIII, Art. 2; pp. 4-72, 21 text figure
ee Memoranda Upon the ‘Anatomy. of. the. Respiratory Tract, -Foregu ;

SO Ls - ‘Thoraeie Viscera of a Fetal Kogia breviceps. “By John D. ‘Kernan, J It.)
ae ee a and HL, yon Ww. Schulte, 1918, ee XXXVUL, aS 8 pp 231-207, |
ag ee Bae "16 text figures. :
ie Be The Skull of Ziphius cavirostris. By John D. Kerman, 1918, : Bulleti ae
ea OS XXXVI, “Art. 11, pp. 349-394, Pls. XXOKEXI.. ipa Z.
. ee The Indigenous Land. Mammals of Porto Rico, Living oad Punk: _ By.

ee ue ie Anthony, 1918, Memoirs, N. Ry a, ee 2, FP 831-436, Pls. LY:
= oe saexiyy Bo GER figures. re Soe
es _Severtzow’ s§ Classification of the Felide.. By J. A Allen, 1919, Bulletin,
J LL Art pp. S35 St =
en . a Notes on the Synonymy and N nena a ‘the: ‘Snialler Spotted Cats f
ee a - Tropical. America. By J. A. Allen, 1919, Soe XLL, ee vp. 8 J
ae Pe 419, 31 text. figures. ae foe Gee
ee oy Mammals Collected i in: Eastern: Cuba i in’ 1917, wa
ee ge ei _ Species. - -By HE Anthony, 1919, Pulse, Xi, Ast. 20, ), PP
ee 643, Pls. XxXV-XXXVE. ee
S ‘New Mammals from eee ; By oe
Legs - Art. 12, pp: 469-475, Pl. “Xxxul, 4 text ene
2 Description of a New Species of Serow. from rene” - Provinee, ¢
eee By Roy Chapman. Andrews, 1921, Novitates, No. 6, pp. 1-3. - 2 os ae
: a ‘Remarkable Case of External Hind Limbs ina Humpback Whale. By mee
ee Pe OY: Chapman | Andrews, 1921, Novitates, No. 9, pp. 1-6. Tees a AG
oo New Mammals from British Guiana oe Colombia... ee H. E. “Anthony, Br rs
: 1921, Novitates, No. 19, pp. AST. aA
3 Preliminary. Report. on Ecuadorean Mammals. ee i. By
RSs atare SAGBAS: Novitates, No, 20, pp. eh ee
Rp reas ~ Preliminary Report on Beuadorean Mammals, -No. 2. ‘By H. EB Anthony, eee =
ce 4922, Novitates, No. 32, pp. 1-7. Se iss
A New Fossil Rodent from Houador. ae H. =, Anthony’,

_ No. ee PP: 1- “4.

H. E “Anthony, |

1922, Novato Z

OLLEDIN A. M. N. HL. Vou. XLVII, Puate V

rawn by Richard Deckert

Idiurus langi J. A. Allen. Medje
(About one-half natural size)

59.9,32(67.5)
Article II—SCIURID#, ANOMALURIDA, AND IDIURIDAi
COLLECTED BY THE AMERICAN MUSEUM CONGO

EXPEDITION!
By THE Late J. A. ALLEN?
PLATE V
CONTENTS

PAGE
(ETc cece elle Aa uA eas ee PG AnD a 39

Species and Subspecies, with their Localities and Number of Specimens
SURE TED, BEyGLS TOPO DAI CESY y R e e iea ae a ecA n te e 40

Localities, Species and Subspecies, and Number of Specimens taken at
ee MeLMe ONAN Re a RS er a he g aperation fdr dhe, OE. SRL E S 42
New Species and Subspecies, with their Type Localities................ 44
PEL EUS), SUITLTOO TE ios ee i UE eee gine Ce Ailes cae a Rear ara Me ee aera ORE! ©
Sciuride...., ere teh tar we Me eS tele) ke hube egg boo ae Sr ga Ct 8 44
EERE PS SEC STOR Dos Oe ed as Re ee a ne 44
Mee TES UE OULMONUUML PUSILOs 2 d.0s0 so c{ecc se ahs Kiet vlage 6a qeininctn es es 47
BASE US TH OUNOEI INE TUCO UGIUUS yin ces i ie ois void a Satan bls e's eile 08 48
NANPOSCLULUS THIOOTACKTUM TUOTLCOLUS.<.. 6.60 cb ec eet eee eee wane cee bees 50
ET LUST TUIVICOION LOLETIS® .¢ ooo. cv cos occ ns waft voclae we hs coe ble ohio e oe O OOL
PEAR ISCU MIS OILENYERTUS: CWETYUNTUS....o roa o's os wale sw celeb ee nels Sga bee's ae ue 51
Me eI SROMY CRUCNTINS AVL > os cc kin ees bk oh bnvera's 4 aides ie «alate gies veh s ae
EAMES NETOMUS!OMGIGO, coc: «ays 0 si v2 1h oleae Lb aed Reve a ebewls els 54
MME SICONMOLCHS (CONQUCUS .. ... cis < de vc bin a's eo Wiehe oie ep eek wile viel ee bo.sinve 56
(DP UGS PRGOILE GUDMIC Ee Oe BRN REET ae a Re ODE eG Pee ger 56
Tamiscus alexandri.......... UY Sch Ae pe sete eee earn Mee RT BS
eRe SESH TN OCKECCHINICOLE: 0 Ste: \odkels Sn = Dae AR See Liens Paneclele we eee s 59
Protoxerus stangeri signatus..... BO ey Ceca ee ad CREM Smeg Svs OT, ok kate eee 61
SECM OTE SIIOMUSIR USS a.) 2 53 x0 Salo ad och ete tole Sen aeons ee |) OL
Anomaluride.... . te BAR Ree Ee eR Cnt ha NG ie Soe ne ae 62
RESINS FOCKSOILE JOCESOM cs <bscc sc a sole ek pd ke ee a wt seve ee senate eT Batt 62
CAD DELTINEINAEL TEOSIINIG MPlee e a 0 rNR 63
EEN LOSI CCONOTLE CRO PUL © 6.2. 5k wipes oie wile See cn we cis eb cae beeline 65
a I ene es nh © 5 on yc Ss eyh, age Pie Sté wd wl wed eitray ales nye wud'ers 68
_ LTE SETPEE 200) EIEN Ae ee ee ee ee 68
AE OTITIS 8b ete e ea ee ae ee a 69
LE TG Tied a Ey Bie alee Oa a rs pe ee 70

INTRODUCTION

The Sciuride, Anomaluride and Idiuride of The American Museum
of Natural History Congo Expedition, collected by Messrs. Herbert
Lang and James P. Chapin during six years (1909-1915) of field work,

‘Scientific Results of the Congo Expedition, Mammalogy, No. 6
2After the author’s demise the manuscript was arranged for Oa heaaee by Herbert Lang.

40 Bulletin American Museum of Natural History [Vol. XLVII

number 480 specimens, representing 20 forms (16 species and 4 additional
subspecies). The collection consists of many well-prepared skins with
skulls, a number of skeletons, and a few specimens preserved in alcohol.
The 315 specimens of Sciuride are referred to 14 forms (10 species and
four additional subspecies), of which three subspecies are described as
new. The 125 specimens of Anomaluride represent three genera and are
referable to 3 forms, one of them new to science. The Idiuride are
represented by 40 specimens of three quite different forms, two of which
are here for the first time described. These far exceed the total number
previously extant in all of the museums of the world, their nocturnal
habits and secretive mode of life rendering their capture difficult.

This is a very valuable accession as The American Museum of
Natural History had but little African material of this order previous to
the reception of the Lang-Chapin Collection. Other museums of this
country have generously placed at my disposal material for purposes of
comparison. To Mr. Gerrit 8. Miller, Jr., Curator of mammals in the
United States National Museum, to Mr. Samuel Henshaw, Director,
and to Dr. G. M. Allen, Curator of mammals, of the Museum of Com-
parative Zodlogy at Harvard University, I am indebted for the loan
of many of their specimens.

The forms, and the number of specimens of each and their localities,
are given in the subjoined lists. :

SPECIES AND SUBSPECIES, WITH THEIR LOCALITIES AND NUMBER OF
SPECIMENS FROM EACH LOCALITY

Speci-
Species and Subspecies Localities mens
Sciuridee

1. Athosciurus poensis (A. Smith) Medje 2, Niapu 1 3
2. Heltosciurus rufobrachium pasha

(Schwann) Faradje 2, Niangara 10 12
3. Heliosciurus rufobrachium medji-

anus, new subspecies Akenge 1, Avakubi 1, Bosobangi 1,

Gamangui 1, Medje 23, Niapu9 36
4. Heliosciurus rufobrachium rubri-

catus, new Subspecies Avakubi 7, Bafwasende 1, Lubila 1 9
5. Heliosciurus multicolor lateris
Thomas Aba 1 1
6. Funisciurus anerythrus anery-
thrus (Thomas) Avakubi 7, Gamangui 6, Medje 8,
Ngayu 2, Niangara 5 28

7. Funisciurus anerythrus niapu, new
subspecies Niapu 22, Stanleyville 2 24

1922] Allen, Congo Sciuride, Ancmaluride, Idiuride 41

Speci-
Species and Subspecies . Localities , mens
8. Funisciurus pyrropus akka de
Winton Akenge 3, Avakubi 1, Boyulu 1,
Gamangui 4, Medje 7, Niangara
3, Niapu 13 32
9. Funisciurus congicus congicus
(Kuhl) Leopoldville 1 i
10. Tamiscus eminit emini (Stuhl-
mann) Avakubi 6, Bafwabaka 7, Batama 1,

Faradje 2, Gamangui 1, Medje
10, Ngayu 4, Niangara 5, Niapu
12, Pawa 1, Poko 1, Stanley-

ville 13 63
11. Tamiscus alexandri (Thomas and
Wroughton) Avakubi 4, Faradje 3, Gamangui 5,
Medje 2, Nala 1, Ngayu 2, Pawa
1, Rungu 1 19

12. Protoxerus stangeri centricola
(Thomas) Akenge 5, Avakubi 5, Bafwabaka 2,
: Faradje 1,Gamangui 4, Kamu-
nionge 1, Medje 6, Ngayu 7,
Niangara 2, Niapu 20, Stan-

leyville 1 54
13. Protoxerus stangeri signatus
Thomas Bolobo 1 1
14. Euxerus erythopus lacustris
(Thomas) _ Faradje 21, Niangara 10, Rungu 1 32
Anomaluridse

15. Anomalurus jacksoni jacksoni de
Winton Akenge 7, Avakubi 2, Gamangui 1,
Medje 28, Niapu 16, Panga 4 58
16. Anomalurella pusilla (Thomas) Akenge 4, Avakubi 1, Medje 36,

Ngayu 2, Niapu 10 5d
17. Anomalurops beecrofti chapini, new
subspecies Akenge 1, Medje 12, Poko 1 14
Idiuridz
18. Idiurus zenkeri zenkeri Matschie Avakubil, Medje 27, Niapu 2 30
19. Idiurus langi, new species Medje 6 6

20, Idiurus panga, new species Panga 4 4

Locaities, SPECIES AND SUBSPECIES, AND NUMBER OF SPECIMENS

Localities

Aba
Akenge

Bafwabaka
Bafwasende
Batama
Bolobo
Bosobangi
Boyulu
Faradje

Gamangui

Kamunionge
Leopoldville
Lubila
Medje

ce
ce

Bulletin American Museum of Natural History

TAKEN AT EACH LOCALITY

Species and Subspecies

Heliosciurus multicolor lateris Thomas

Heliosciurus rufobrachium medjianus, new subsp.

Funisciurus pyrropus akka de Winton
Protoxerus stangeri centricola (Thomas)
Anomalurus jacksoni jacksoni de Winton
Anomalurella pusilla (Thomas)
Anomalurops beecroftt chapini, new subsp.

Heliosciurus rufobrachium medjianus, new subsp.
Heliosciurus rufobrachium rubricatus, new subsp.

Funisciurus anerythrus anerythrus (Thomas)
Funisciurus pyrropus akka de Winton

Tamiscus emini emini (Stuhlmann)

Tamiscus alexandri (Thomas and Wroughton)
Protoxerus stangeri centricola (Thomas)
Anomalurus jacksoni jacksoni de Winton
Anomalurella pusilla (Thomas)

Idiurus zenkeri zenkeri Matschie

Tamiscus emini emini (Stuhlmann)

Protoxerus stangeri centricola (Thomas)
Heliosciurus rufobrachium rubricatus, new subsp.
Tamiscus emini emini (Stuhlmann)

Protoxerus stangeri signatus Thomas
Heliosciurus rufobrachium medjianus, new subsp.
Funisciurus pyrropus akka de Winton
Heliosciurus rufobrachium pasha (Schwann)
Tamiscus emini emini (Stuhlmann)

Tamiscus alexandri (Thomas and Wroughton)
Protoxerus stangert centricola (Thomas)

Eucerus erythopus lacustris (Thomas)
Heliosciurus rufobrachium medjianus, new subsp.
Funisciurus anerythrus anerythrus (Thomas)
Funisciurus pyrropus akka de Winton

Tamiscus emini emini (Stuhlmann)

Tamiscus alexandri (Thomas and Wroughton)
Protoxerus stangeri centricola (Thomas)
Anomalurus jacksonijacksoni de Winton
Protoxerus stangeri centricola (Thomas)
Funisciurus congicus congicus (Kuhl)
Heliosciurus rufobrachium rubricatus, new subsp.
Atthosciurus poensis (A. Smith)

Heliosciurus rufobrachium medjianus, new subsp.
Funisciurus anerythrus anerythrus (Thomas)
Funisciurus pyrropus akka de Winton

BPP WNN RPE RE EP NON RFP RE NOR OEP NN RP RP BRNO wee

bo bo
OW HE HS ROH RO

“I CO

[Vol. XLVII

Totais

Se ee

29:

1922]

Localities
Medje
“é

Niangara
(<4

Panga

Pawa
74

Poko

Rungu

“ec

Stanleyville

Allen, Congo Sciuride, Anomaluride, [diuride

Species and Subspecies

Tamiscus emini emini (Stuhlmann)

Tamiscus alexandri (Thomas and Wroughton)
Protoxerus stangert centricola (Thomas)
Anomalurus jacksoni jacksoni de Winton
Anomalurella pusilla (Thomas)

Anomalurops beecrofti chapini, new subsp.
Idiurus zenkeri zenkeri Matschie

Idiurus langi, new sp.

Tamiscus alexandri (Thomas and Wroughton)
Funisciurus anerythrus anerythrus (Thomas)
Tamiscus emini emini (Stuhlmann)

Tamiscus alexandri (Thomas and Wroughton)
Protoxerus stangeri centricola (Thomas)
Anomalurella pusilla (Thomas)

Heliosciurus rufobrachium pasha (Schwann)
Funisciurus anerythrus anerythrus (Thomas)
Funisciurus pyrropus akka de Winton
Tamiscus emini emini (Stuhlmann)

Protoxerus stangeri centricola (Thomas)

- Euxerus erythopus lacustris (Thomas)

Athosciurus poensis (A. Smith)

Heliosciurus rufobrachium medjianus, new subsp.

Funisciurus anerythrus niapu, new subsp.
Funisciurus pyrropus akka de Winton
Tamiscus emini emini (Stuhlmann)
Protoxerus stangert centricola (Thomas)
Anomalurus jacksoni jacksoni de Winton
Anomalurella pusilla (Thomas)

Idiurus zenkeri zenkeri Matschie
Anomalurus jacksoni jacksoni de Winton
Idiurus panga, new sp.

Tamiscus emini emini (Stuhlmann)
Tamiscus alexandri (Thomas and Wroughton)

. Tamiscus emini emini (Stuhlmann)

Anomalurops beecrofti chapini, new subsp.
Tamiscus alexandri (Thomas and Wroughton)
Euzxerus erythopus lacustris (Thomas)
Funisciurus anerythrus niapu, new subsp.
Tamiscus emini emini (Stuhlmann)
Protoxerus stangeri centricola (Thomas)

ie) = =
NOOF- ONOAWHONN NH FN KH O&O

he Re
On w

— et
Ee dD © O

=
BH WoO Fe Bee Be

Totals

167

Bt)

105

44 Bulletin American Museum of Natural History [Vol. XLVII

New SPECIES AND SUBSPECIES, WITH THEIR Type LOCALITIES

1. Heliosciurus rufobrachium medjianus. Medje
2. Heliosciurus rufobrachium rubricatus. Lubila
3. Funisciurus anerythrus niapu. Niapu
4. Anomalurops beecroftt chapini. Medje
5. Idiurus langi. Medje-
6. Idiurus panga. Panga
GENERAL SUMMARY
Species and
Families Genera Subspecies Specimens Localities!
Sciuridee 6 14 315 23
Anomaluridze 3 3 125 8
Idiuridee eo 3 40 4
10 20 480
SCIURIDE

ZETHOSCcIURUS Thomas

Asthosciurus THomas, 1916, Ann. Mag. Nat. Hist., (8) XVII, March, p. 271.
Genotype, by original designation, Sciurus poensis A. Smith.

Atthosciurus (subgenus of Heliosciturus) Houuister, 1919, U. S. Nat. Mus.
Bull. 99, part 2, May 16, p. 9.

AEthosciurus poensis (A. Smith)

Sciurus poensis A. SmitH, 1835, South African Quart. Journ., II, p. 64. Fernando
Po (Gray).

Aithosciurus poensis THomas, 1916, Ann. Mag. Nat. Hist., (8) XVII, p. 271.

Three specimens: Medje, 2 (c’ and 9 adult), January 24, 1909;
Niapu, 1 (co adult), November 28, 1913.

Collectors’ measurements of the Medje specimens: Total length,
o 322 mm., 2 337; head and body, o 144, Q 152; tail vertebra, 7
178,°9 185; bind feot,.c! 35,2" 35;-ear.o 1470.14.

Skull, same specimens: Greatest length, co 37.3, 2 38.2; zygomatic
breadth, not measurable.

The Niapu specimen is a little smaller and less iatieee

Entire pelage, including feet and tail, olivaceous gray, except ventral
surface, which is washed with ochraceous medially, the color of the sides
extending over the lateral third of the ventral area from axille to loins.

Compared with two specimens of #thosciurus poensis (No. 8639,
Kribi, Cameroon, and No. 15667, Mus. Comp. Zodl., Lolodortf,
Cameroon), with which they closely agree. Larger series from the two
regions (Cameroon coast and Upper Congo) might indicate an appreci-
able average difference not indicated by the material now available.

1The total number of localities at which these forms were collected is 24.

1922] Allen, Congo Sciuride, Anomaluride, Idiuride 45

Hetiosciurus Trouessart

Heliosciurus (subgenus of Sciurus) Trouessart, 1880, Le Naturaliste, II,
October, p. 292. Genotype, by subsequent designation (Thomas, 1909), Sciwrus
gambianus Ogilby. Trouessart originally designated Sciurus annulatus Desmarest as
the type, but as this species is not positively identifiable Thomas has, with good
reason, replaced it by Sciurus gambianus Ogilby.

The Heliosciurus rufobrachium' group is represented in the Lang-
Chapin Collection by 57 specimens, collected in the region comprised
between Avakubi and Bafwasende, south of the Ituri-Aruwimi River,
northward to Niangara, on the Uele River, and eastward to Faradje.
Three geographical areas are thus included—(1) the region south of the
Ituri-Aruwimi covered with Rain Forest, (2) the forested area between
the Ituri-Aruwimi and Bomokandi-Uele rivers, and (3) the Uele bush-
veldt district to the north. The specimens from these districts, when
arranged serially, show well-marked differences in coloration in correla-
tion with the varying conditions of the districts, the extremes represented
—the Avakubi specimens on the one hand and the Niangara-Faradje
specimens on the other—being so widely different that, without the con-
necting series from intermediate points, they might readily be con-
sidered as possibly specifically separable, especially if represented by a
single specimen or even by a small series of specimens. The specimens
from the intermediate localities show, however, unmistakable inter-
gradation between the two extreme geographic phases. The differences
are primarily the amount and intensity of rufous on both fore and hind
limbs and the color of the whole ventral area, particularly of the throat
and inside of the limbs. Also the northern pale veldt form is distinctly
smaller than either of the two forest forms. The main feature of differ-
entiation is the steadily increasing erythrism of the ventral surface and
limbs from the northern veldt country to the heavy Rain Forest south of
the Ituri.

In the coloration of the upper surface there is little to distinguish
the specimens from the three areas, although the Niangara and Faradje
specimens average somewhat paler than those from farther south.

1Mr. Lang has called my attention to the fact that Sciurus rufobrachium Waterhouse has nearly
three months priority over S. rufobrachiatus of the same author and it should therefore be accepted
in place of the latter. The species so long universally known as Sciurus rufobrachiatus was named
Sciurus rufobrachium by Waterhouse in an incidental reference to it in a footnote to a paper in the
‘Annals and Magazine of Natural History’ (X, p. 202) published in November 1842, but in his formal
description of the species (Proc. Zoé]. Soc. London for 1842, p. 128, published January 1843) he changed
the name to Sciurus rufobrachiatus, without explanation or reference to the earlier name for the same
species. Consequently the name Sciurus rufobrachium was used by just a few, Fitzinger (1867) being the
last. All succeeding authors have either overlooked or ignored the earlier name till it was brought to
light by Mr. Lang in the preparation of his paper on the bibliography and distribution of African
members of this group.

It may be noted alsothat Waterhouse’s Sciurusleucogenys has priority over his Sciurus erythrogenys,
the two cases being parallel and dating from the same footnote.

46 Bulletin American Museum of Natural History [Vol. XLVII

Through individual variation in the general tone, specimens from either
of the series can be selected which are mutually indistinguishable in the
color of the upperparts. It is quite different, however, with the ventral
surface. In the northern form (H. 7. pasha) it is pale yellowish white,
varying in different specimens from dull whitish to faintly yellowish on
the median area from the throat to the anal region, with usually a large
whiter pectoral area, the sides being darkened by the dark basal portion
of the hairs showing through the superficial light tipping. The chin and
throat are a little browner than the foreneck and breast, being sometimes
dull yellowish brown, but rarely approaching rufous. The median
portion of the extreme base of the under side of the tail has sometimes
a slight rufous tone. The outer edge of the forearm and the upper surface
of the manus vary from dark rufous to brownish rufous, but the inside of
the forearm is pale like the ventral surface, usually without trace of
rufous. |

In specimens of the other extreme, taken south of the Ituri near
Avakubi, the whole lower surface of the body has a strong rufous tone,
the throat, foreneck, sides of breast, inguinal region, and entire inner
surface of both fore and hind limbs are intense vivid rufous, as is also a
conspicuous median patch at the base of the under side of the tail, and
the upper surface of the fore and hind feet are also red. The median
ventral area, from the chest to the lower abdomen, is pale rufous grizzled
slightly with black.

The three forms may be characterized as follows.

1. Light northern form: Underparts superficially pale, the hair-tips whitish or pale
yellowish, usually a rather distinct narrow median light band (often broad-
ening at pectoral region) contrasting with a much darker and broader area on
either side from axille to loins; throat and inside of limbs light, uniform in
color with the central light portion of ventral surface, except wrists; out-
side of hind limbs like back; outer edge of forearm and upper surface of feet
pale rufous. On hind limbs the brownish-rufous tone is usually restricted to
upper surface of feet; in exceptionally erythric specimens it may extend to
the lower leg and include the inner surface as well as the outer, thus forming a
dull rufous band just above the ankle, and even extend up the inner side of
the leg, with a similar extension of rufous on the inner surface of the lower
HORE AMM aes 8 pete AEs Mose Oe nope cued tee Mae CROANIE hesat oeerte H.r. pasha (Schwann).

2. Darker middle form: Underparts darker, nearly uniform except for a small
sharply defined white pectoral area, the hairs ringed basally with black and
buff-tipped; throat and inside of limbs pale or dull rufous, in contrast with
abdominal region; outer surface of hind limbs like back; outer edge of fore
limbs intense rufous, which encircles the lower forearms and lower legs and
includes the upper surface of fore and hind feet.

H. r. medjianus, new subspecies.

1922] Allen, Congo Sciuride, Anomaluridx, Idiuride 47

3. Darker southern form: Underparts medially strongly suffused with pale rufous,
usually without trace of a whitish pectoral area; throat, sides of head below
eyes, entire inner side of fore and hind limbs, and extreme posterior part of
abdomen intense dark rufous; outer edge of fore limb, lower forearm in front
and upper surface of fore and hind feet, wrists and ankles chestnut-rufous,
which also extends to the median basal underside of tail.

H. r. rubricatus, new subspecies.

Average External and Cranial Measurements of Heliosczurus
rufobrachium pasha, medjianus, and rubricatus

External | Cranial

| | | | Wecehecs

| | ! Pa | | | = | 5

lap etteal tema ee | a ipoe

Se ed = ic | open 2 =

a S = 5 = Z 3

ae = = = sah a & I

ees g 5 Bet hile g 5:

Zn | & ic = mo | 8 G w
H.r. pasha, Niangara | 10 | 473 | 223 | 251 | 56.2 | 18.3| 52.2 | 30.6
H.r. medji- Medje and 12 497 | 239 | 258 | 58.2 | 18.0 | 52.9 | 30.5
anus, Niapu eG 507 | 239 | 268 | 58.3 | 17.5 | 53.4 | 30.8
H.r. rubricatus, Avakubi, 6 | 501 | 234 262 | 59.3 | 18.3 | 53.4 | 31.9

The relation of these three forms is so obviously geographic, and
reflects so strikingly the effect of environment, it seems desirable to
recognize these facts nomenclaturally, as follows.

| Heliosciurus rufobrachium pasha (Schwann)
_Sciurus rufobrachiatus pasha Scuwann, 1904, Ann. Mag. Nat. Hist., (7) XIII,
January, p. 72. Type locality, Bellima, Mombuttu, Belgian Congo.

Heliosciurus rufobrachiatus pasha Tuomas, 1915, Ann. Mag. Nat. Hist., (8)
XVI, December, p. 473, (part). The Irumu specimen.

Represented by 12 specimens (7 males, 5 females), all adult, col-
lected as follows:

Niangara, 10 (6 o&, 4 2), November 9-20, 1910.

Faradje, 2 (o’, 2), December 2, 1911.

The type locality of Sciurus rufobrachium pasha Schwann is given
as “Bellima, Mombuttu,” the type being an adult male collected by
Emin Pasha, July 13, 1883. Bellima does not now exist, Mr. Lang in-
forms me. It was long since abandoned by the natives. But its former
site was about 25 miles southeast of the present Niangara. Hence the
ten specimens in the Lang-Chapin Collection are practically topotypes,
from which the two from Faradje are indistinguishable.

48 Bulletin American Museum of Natural History [Vol. XLVII

Collectors’ measurements of the Niangara series (6 males, 4 females) :

Total Length Head and Body Tail Vertebre Hind Foot Ear
o& 478 (470-488) 223 (216-235) 250 (245-260) 57.3 (54-59) 18.7 (17-19)

Q 467 (455-479) 217 (215-227) 250 (242-257) 55.0 (53-56) 18.5 (17-20)
Skulls, same specimens:
Greatest Length Zygomatic Breadth
(=occipito-nasal length) (=greatest breadth)
oo 52.4 (51.0-54.0) 30.3 (29.1-31.3)
9. db1.7 (9.2-55.7) 30.8 (28.8-32.0)

This form is readily distinguishable from those recorded below from
localities more to the southward, including Medje, Niapu, and Avakubi,
by its much paler general coloration and very much lighter underparts.

Heliosciurus rufobrachium medijianus, new subspecies

Heliosciurus rufobrachiatus pasha THomas, 1915, Ann. Mag. Nat. Hist., (8)
XVI, December, p. 473, (part). The Medje and ‘‘Poko”’ specimens. ‘‘ Poko” =
south of Poko, in forest probably nearer Niapu than Poko.

Type, No. 50761, @ adult, Medje, Belgian Congo, January 17, 1910; Herbert
Lang and James P. Chapin. Orig. No. 487.

Larger and darker than H. 7. pasha, the underparts very much darker, the white
tips of tail hairs much shorter, inside of limbs with much more rufous. Differs from
H.r. rubricatus in lacking the strong rufous suffusion of the underparts and the intense
rufous of inside of limbs, throat and anal region.

Collectors’ measurements of type: Total length, 502 mm.; head and body, 224;
tail vertebree, 278; hind foot, 59; ear, 20.

Skull (type): Greatest length (=occipito-nasal length), 53.6; condyloincisive
length, 49.3; least interorbital breadth, 15.8; tip to tip of postorbital processes, 23.7;
postorbital breadth, 16.1; breadth of braincase, 22.2; zygomatic breadth, 30.7;
length of nasals, 16.1; breadth of nasals anteriorly, 8.4, do. at posterior border, 5;
length of maxillary toothrow, 10. .

Represented by 36 specimens (of which 23 are topotypes), taken as follows:

Avakubi, north side of Ituri River toward Bosobangi, 1 (“ adult), April 11, 1914.

Bosobangi, 1 ( 9 adult), December 24, 1909.

Gamangul, 1 (o adult), February 7, 1910.

Medje, 23 (13 adult, 10 immature): 9 (6.7, 3 9, all adult), January 15-20, 1910;
10 (50%, 5, all immature, ranging in age from nurslings to half-grown), April 10,
August 3-September 14, 1910; 4 (207, 29, adults—1 complete skeleton), March
22, 1910; February 28, April 2, 6, 1914.

Niapu, 9 (7o7, 29, of which 6 are adult and 3 immature), November 14—
December 8, 1913.

Akenge, 1 ( 9 adult), October 16, 1913.

Collectors’ measurements of 13 adults (8 «, 5 9) from Medje:

Total Length Head and Body Tail Vertebrz Hind Foot Ear
o& 497 (441-534) 241 (225-254) 256 (211-292) 58.4 (55-62) 18.5 (17-20)
Q 495 (475-502) 231 (224-238) 264 (246-278) 57.5 (56-59) 18.2 (17-20)

1922] Allen, Congo Sciuride, Anomaluridx, Idiuridxe 49

Skulls, same specimens:

Greatest Length Zygomatic Breadth
guy 00.0 (0) 7—54. 9) 30.7 (28.9-31.9)
@ 52.5 (51.4-53.6) 30.5 (29.4-31.6)

Collectors’ measurements of 6 adult specimens (407, 2 9) from Niapu:

Total length, 507 (487-524); head and body, 239 (229-247); tail vertebre
268 (255-280); hind foot, 58.3 (53-61); ear, 17.5 (17-18).

Skull (5 of the same specimens, 4 o’, 1 9): Greatest length, 53.4 (52.4—54.2);
zygomatic breadth, 31.0 (29.8-31.6).

Collectors’ measurements of 4 specimens (2 o’, 2 2) from other localities near
Niapu (Bosobangi 2, Gamangui 1, Akenge 1):

Total length, 493 (452-542); head and body, 237 (225-251); tail vertebre, 265
(248-291); hind foot, 54.5 (53-56); ear, 18 (16-19).

Skull (same specimens): Greatest length, 52.8 (51.9-53.9); zygomatic breadth,
50:3 (29.7-31.8).

The specimens from Niapu and other localities near Niapu agree
closely in coloration and other features with the type series from Medje.
The large series from Medje is especially interesting from the fact that
it contains a large number of young specimens, ranging in age from
nurslings to nearly half grown. Of six nurslings (the only teeth pre-
sent are the incisors) one was taken April 10, and five August 3, 5,
and 24, three of them being from the same litter. Four others are a
week or two older (taken September 2, 9, 18, 14) with the first cheek-
teeth just breaking through the gums. The coloration of these young
specimens differs from that of adults of the same series in no mate-
rial respect in either pattern or color. The pelage is, of course,
much softer with more underfur, and the ventral surface is more heavily
clothed, and hence the color areas are more sharply outlined. The hair of
the dorsal surface differs from that of adults in the annulations being
apparently narrower, an effect due probably to the shorter pelage. The
hairs on the sides of the body are minutely tipped with whitish passing
gradually into buffy or pale fulvous toward the midline of the back, as is
the case with adults, so that the surface effect is exactly as in the latter.
On the ventral surface the color areas are more sharply defined than in
adults, owing to the thinner and less complete cqndition of the coat in the
latter. All have the pectoral white patch indicated, and in nearly all
it is pure white and forms a conspicuous mark, although varying greatly
in size in different individuals. Those in which it is largest have also a
small tuft of pure white soft hairs at the axillz, which is obsolete in those
that have the pectoral mark only slightly developed. The sides of the
nose, chin, and upper throat are dull yellowish brown, becoming paler
posteriorly. The inside of the fore and hind limbs is pale yellowish rufous,

50 Bulletin American Museum of Natural History [Vol. XLVII

which deepens on the lower hind limbs and anal region to a much darker
tone, and is more or less strongly diffused over the basal portion of the
underside of the tail. The mid-abdominal region (axillz to loins) is a
grizzle of dull brown and pale buffy, with a tendency to a lighter median
line. The upper surface of the fore and hind feet is mixed dark rufous
and black, the black basal portion of the hairs showing more or less at the
surface.

Individual color variation in adults is due primarily to the amount of
rufous suffusion present, varying from a strong rufous tone throughout
the pelage to its almost entire absence. Specimens of either of these
types, however, are exceptional. The specimen selected as type of
medjianus represents the average condition. The extreme rufous ex-
amples strongly approach rubricatus and indicate intergradation between
the two forms. The specimens recorded by Thomas from Medje and
Poko as referable to H. r. pasha (loc. cit.) should doubtless be referred to
medjianus, since these localities are in the type region of the latter.

Heliosciurus rufobrachium rubricatus, new subspecies

Type, No. 50748, o adult, near the Lubila River, an affluent of the Tshopo River,
about 50 miles southwest of Avakubi (south of the Ituri River), Belgian Congo,
September 20, 1909; Herbert Lang and James P. Chapin, Orig. No. 123.

Similar to H. r. medjianus in size and color of upperparts; underparts more
strongly suffused with rufous; inside of fore and hind limbs and anal region intense
dark rufous; upper surface of feet, wrists and ankles, and median basal underside
of tail chestnut-rufous. .

Collectors’ measurements of type: Total length, 552 mm.; head and body, 255;
tail vertebre, 297; hind foot, 61; ear, 19.

Skull (type): Greatest length (=occipito-nasal), 55.4; condyloincisive length,
51.2; least interorbital breadth, 16.7; tip to tip of postorbital processes, 25.6;
postorbital breadth, 14.4; breadth of brain-case, 23.1; zygomatic breadth, 31.5;
length of nasals, 17.6; breadth of nasals anteriorly, 8.3, do. at posterior border, 6.6;
length of maxillary toothrow, 10.9.

Represented by 9 specimens, as follows:

Avakubi, 7 (3 o, 3 9, all adult, 1 in alcohol), October 1, 13, December 8, 1909,
January 12, 24, June 22, and August 26, 1914.

Bafwasende (35 miles south of Avakubi), 1 (o adult), September 23, 1909.

Lubila, 1 (o adult), September 20, 1909.

Collectors’ measurements of 6 specimens (3 o, 3 @) from Avakubi: Total
length, 501 (482-525); head and body, 234 (226-253); tail vertebree, 262 (251-279);
hind foot, 59.3 (54-63); ear, 18.3 (17-20).

Skull (4 of same specimens—2 too much broken for measurement): Greatest
length, 53.4 (52.4-54.8); zygomatic breadth, 31.9 (29.7-83.4).

The relation of the present form to medjianus has been indicated in
the detailed comparison already given (pp. 45 to 47) of the three forms

1922] Allen, Congo Sciuride, Anomaluride, Idiuride Dik

of the rufobrachium group represented in the present collection. The
differences that distinguish medjzanus from pasha are greatly intensified
in rubricatus, the northward range of which appears to be limited by the
Ituri River.

Heliosciurus multicolor Group
Heliosciurus multicolor lateris Thomas

Heliosciurus multicolor lateris THomas, 1909, Ann. Mag. Nat. Hist., (8) IV,
August, p. 102. Type locality, Lado, Mongalla.

Represented by one specimen, subadult female, Aba, Belgian Congo,
December 12, 1911.

Collectors’ measurements: Total length, 390 mm.; head and body,
185; tail vertebree, 205; hind foot, 45 (s. u. 42.5); ear, 15.

Skull: Greatest length, 45.4; condyloincisive length, 40.7; zygo-
matic breadth, 36.

Provisionally referred to this subspecies, with the description of which
it well agrees.

Funisciurus Trouessart

Funisciurus (subgenus of Sciurus) TrovEssartT, 1880, Le Naturaliste, II, No.
37, October 1, p. 298. Genotype, by monotypy, Sciurus isabella Gray = Funisciurus
lemniscatus isabella (Gray). A few weeks later (idem, 1880, II, No. 40, November 15,
p. 315) he designated Sciurus lemniscatus LeConte as type of Funisciurus, on the
assumption that S. isabella Gray was a strict synonym of the earlier S. lemniscatus.

Funisciurus anerythrus anerythrus (Thomas)

Sciurus pyrrhopus anerythrus THomas, 1890, Proc. Zo6l. Soc. London, pp. 447,
448, PI. XL, animal. Two specimens. Type locality, Buguera.

Funisciurus anerythrus THomAs, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 473. Mawambi (1), Avakubi (1), Medje (6), Poko (15 specimens).

Represented by 28 specimens, taken at five localities, as follows:

Avakubi, 7 (2 o’, 1 adult, 1 immature; 5 9, all adult), October 7,
November 5, 24, 1909, February 11, April 8, 1914.

Ngayu, 2 (2 9—1 adult, 1 immature), December 17, 1909.

Gamangui, 6 (1 o’, 5 9—2 2 immature), January 30, February 7,
15, 19, 1910.

Medje, 8 (38 o', 5 9—2 oO, 2 GY immature), January 18-24,
September 26, October 5, 1910, April 3, May 25, June 15, 1914.

Niangara, 5 (1 o adult, 4 9 —2 immature), November 7, 15, 20, 1910.

Collectors’ measurements of 14 adult specimens (4 males, 10 females,
all middle-aged to adult) from Ngayu, 1 (9); Avakubi, 4 (1 o&, 3 @);
Gamangul, 4 (1 o’,3 2); Medje, 2 (1%, 1 2); Niangara, 3 (10,2 9).

52 Bulletin American Museum of Natural History [Vol. XLVII

Total Length Head and Body Tail Vertebre Hind Foot Ear
355 (346-368) 186 (183-193) 165 (163-175) 46.5 (46-48) 18.0 (17-19)
Q 356 (348-365) 189 (181-195) 168 (160-183) 47.1 (45-49) 17.4 (16-19)

The above measurements indicate a slightly greater size for the
females, as is the case also in F’. pyrropus akka from about the same locali-
ties.

The underparts vary considerably in the amount of buffy or ochrace-
ous wash on the ventral surface, independently of season or locality, the
palest being nearly white and a few (four out of twenty-eight) closely
approaching the average of the Niapu series of twenty-two described
below.

Funisciurus anerythrus niapu, new subspecies

Type, No. 50877, & adult, Niapu, Belgian Congo, November 9, 1913; Herbert
Lang and James P. Chapin, Orig. No. 2120.

Similar to typical anerythrus, but underparts averaging much darker and more
ochraceous, the ochraceous tips of the hairs being longer and brownish ochraceous,
often wholly concealing the dark basal portion of the pelage.

Collectors’ measurements of the type: Total length, 359 mm.; head and body,
190; tail vertebre, 169; hind foot, 47; ear, 18.

Skull (type): Greatest length (=occipito-nasal), 45.7; condyloincisive length,
40.7; least interorbital breadth, 11.7; tip to tip of postorbital processes, 19.2; post-
orbital breadth, 16.1; breadth of brain-case, 20.1; zygomatic breadth, 25.5; length
of nasals, 12.2; breadth of nasals anteriorly, 4.6, do. posteriorly, 5.9; length of
maxillary toothrow, 8.1.

Represented by 24 specimens collected as follows:

Niapu, 22 (10 «1, 12 92—3 & and 4 @ immature) all collected November 9-30,
1913, except one taken at same locality a month later (December 25).

Stanleyville, 2 (o, 92 in alcohol), September 6, 1909, January 18, 1915.

Collectors’ measurements of 13 adult specimens from Niapu (6 males, all adult;
7 females, of which 3 are young adults) :

Total Length Head and Body Tail Vertebre Hind Foot Ear
& 356 (337-390) 195 (181-226) 162 (155-169) 45.7 (45-47) 17.3 (17-19)
Q@ 350 (340-364) 199 (178-205) 163 (157-165) 46.0 (45-47) 17.3 (16-18)

Skulls, same specimens:

Greatest Length Zygomatic Breadth
o 46.5 (48.6-49.0) 25.7 (24.7-26.7)
9 46.9 (44.9-49.7) 25.2 (24.4-26.3)

It is clearly evident that the marked difference in the coloration of
the underparts, which alone distinguishes strongly the Niapu specimens
from all of those from the other six localities to the eastward, is not
seasonal rather than geographic, the fact being that the greater part of the
adults from the other localities were taken at the same season (most of
them during the same month) as those from Niapu. A series of five from

1922] Allen, Congo Sciuride, Anomaluride, Idiuride 53

Niangara were all taken in November; five others from Avakubi were
taken October 7 to November 24; five from Medje were also taken
near the same season (September 26, October 5, and January 18-24).
When laid out in two series, the specimens from Niapu in one and
those from the other localities in the other, it is seen that only a few
of the most heavily colored specimens from the eastern localities equal
the palest of the Niapu series in either the extent or intensity of the
ochraceous wash of the ventral surface. While a few of the lighter
colored specimens in the Niapu series can be matched by a few of
the darkest specimens in the other, and thus indicate intergradation, the
average difference is striking, particularly when the palest specimens of
the two series are compared. It seems desirable therefore to recognize the
Niapu series in nomenclature as a saturate type of the group, especially
since a pale form of anerythrus (F. a. bandarum Thomas) has been desig-
nated from the upper Shari River.

In the present connection it may be of interest to give the results of a
comparison, especially in respect to size, of the F. pyrropus akka series
of thirty-two specimens with the fifty-two of the F. anerythrus group,
since both were collected at the same time at about the same localities.
First it may be stated that the two forms of anerythrus show no differ-
ence in size, and the same is true also of the akka and anerythrus series, in
either external or cranial measurements. Nor am I able to distinguish
the skull of akka by any feature from the skull of anerythrus. In the
coloration of the upperparts there is also a close resemblance, the chief
distinction being the color of the lateral line, which is pale buffy in akka
and white in anerythrus, often indistinct in both. The tail is also alike
in both, on both surfaces. But the difference in the color of the outside
of the limbs (including the upper surface of the feet) and the ventral
surface is striking. In anerythrus the legs and feet externally and the
sides and front of the head are brown with a dull cinnamon-rufous suffu-
sion; in akka intense brownish rufous, particularly on the hind limbs and
feet. In anerythrus the whole ventral surface and inside of limbs is
heavily washed with ochraceous (ochre-yellow to ochraceous rufous),
the basal third of the hairs pale plumbeous; in akka everywhere clear
white to the base of the hairs. As both anerythrus and akka live together
abundantly at all localities from which either is represented in the

__?The Niangara series, as might be expected, is the palest of all, but the Avakubi specimens merge
with them; the palest specimen is an old male from Avakubi, taken October 7, which is white below
with a slight buffy wash over the thoracic region.

54 Bulletin American Museum of Natural History [Vol. XLVII

present collection, with not a single intermediate in a joint series of
eighty-four specimens, it is evident that their status is that of distinct
species. Yet in measurements and proportions and in the coloration of
the upperparts (front and sides of head and lateral line excluded) the
two forms are practically indistinguishable. In respect to measurements,
two series of adults, comprising 13 specimens of nzapu and 14 of akka,
afford the following data: |

External Measurements

Total Head and Tail Verte- Hind Foot Ear

Length Body bree
F. anerythrus niapu = 345 184 161 45.9 17.4
F. pyrropus akka 345 186 159 46.7 17.

Skull Measurements

Greatest Length Zygomatic Breadth
F. anerythrus niapu 47.4 25.2
F. pyrropus akka 46.8 24.2

The measurements and proportions are thus practically identical in
the two forms, the averages in external measurements (taken from speci-
mens in the flesh by the collectors) varying from 0 to 2 mm., and those of
the skulls from 0.6 to 1 mm.

Funisciurus pyrropus akka de Winton

Sciurus pyrrhopus THOMAS, 1888, Proc. Zo6él. Soc. London, p. 9. Two specimens.
Type locality, Tingasi, Monbuttu, Belgian Congo.

Sciurus emini DE WINTON, 1895, Ann. Mag. Nat. Hist., (6) XVI, August, p. 197.
Not Sciurus emini Stuhlmann, 1894.

Funisciurus akka DE WinTON, 1899, Ann. Mag. Nat. Hist., (7) IV, December,
p. 357. To replace Sciurus emini de Winton, preoccupied.

Funisciurus akka Toomas, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 473. Medje (4), Poko (7 specimens).

Represented by 32 specimens, taken at 7 localities, as follows:

Boyulu, 1 ( & adult), September 22, 1909.

Avakubi, 1 (@ adult), July 16, 1914.

Gamangui, 4 (3 & adult, 1 2 immature), January 28, February 7,
18, 1910.

Medje, 7 (2 #7, 5 2—4 immature), January 23, March 25, April 1, 4,
September 4, 29, 1910, June 24, 1914.

Niapu, 13 (5 7, 8 @, all adult), November 12-30, December 27,
AOI ean. |

Akenge, 3 (1 o&, 2 2), September 29, October 10, 11, 1918.

Niangara, 3 (1 0, 2 9, all adult), November 9, 10, 1910.

1922] Allen, Congo Sciuride, Anomaluridx, Idiuride SB)

Collectors’ measurements of 13 youngish adults from Niapu (5
males, 8 females) : :

Total Length Head and Body Tail Vertebre Hind Foot Ear
S&S 387 (317-355) 186 (176-201) 151 (141-159) 47.6 (47-48) 18.2 (18-19)
Q 349 (322-371) 189 (165-206) 160 (145-173) 47.6 (45-49) 17.5 (16-18)

Skulls, same specimens:

Greatest Length Zygomatic Breadth
So 46.2 (45.448.1) 25.5 (24. 7-26. 2)
Q@ 45.7 (45.2-47.9) 25.4 (25.0-26.8)

Only three of the thirteen Niapu specimens of which measurements
are given above are old adults (1 o, 2 9 ), as indicated by the condition
of the teeth and sexual organs (scrotum present in the male, mammez
conspicuous in the females); in the othér ten the dentition is complete
but the teeth are unworn or only slightly worn, and no mamme are
distinguishable in the females. The measurements below of thirteen
specimens from other localities (all that are available) include only
adults of middle age or older, and thus average, as would be expected,
larger than the Niapu series, which consists almost entirely of rather
young adults.

Collectors’ measurements of 13 specimens (6 males, 7 females, all
middle aged or old) from other localities (Akenge, Avakubi, Boyulu,
Ngayu, 1 each; Niangara, 2; Gamangui, 3; Medje, 4):

Total Length Head and Body Tail Vertebre Hind Foot Ear

& 335 (322-351) 190 (184-208) 148 (136-156) 48.5 (48-49) 18.7 (18-20)
Q@ 344 (317-340) 193 (187-202) 151 (180-161) 47.0 (46-48) 18.4 (18-20)

Skulls, same specimens:

Greatest Length Zygomatic Breadth
o 47.5 (46.4-48.5) 25.7 (24.9-27.1)
Q 47.9 (46.4-48.5) 26.3 (24.9-26.7)

The above measurements of two series of specimens, totaling 11
males and 15 females, indicate that the females are slightly larger thar
the males, but the difference is too small to be diagnostic.

The thirty-two specimens of F. p. akka are exceedingly ¢onstant 11
coloration, season and age making very little difference in this respect:
The underparts, including the inside of both fore and hind limbs, aré
pure white to the base of the fur. Immature and September-November
adult specimens show a tendency to a faint pinkish wash on the inside of
the hind limbs and (exceptionally—in two or three speéimeénsg oily) on
the middle of the belly. The pale buff lateral line running from thé
shoulder to the hip varies little in color, but is much better defined in

56 Bulletin American Museum of Natural History [Vol. XLVII

some specimens than in others, the posterior half occasionally becoming
almost obsolete. The dark color of the back extends laterally to a little
below the lateral line, which thus appears to be bordered along the lower
side by a narrow dark band, the flanks being lighter and more suffused
with yellowish than the dorsal area. The rufous of the outside of the
limbs and upper surface of the feet varies considerably in tone, from
light intense rufous to dull brownish rufous. The red on the sides and
front of the head shares this variability of tone.

Fully adult specimens are also very constant in size, the chief varia-
tion being in the length of the tail, which may be somewhat shorter or
longer than the normal length, thus contributing a variable element in
the total length. The tail vertebre are considerably shorter than the
head and body, forming about 46 per cent of the total length, and about
84 per cent of the head and body length. The two pairs of mamme are
both inguinal.

Funisciurus p. akka differs from typical pyrropus in slightly smaller
size and in the rufous of the limbs and head being less vivid and of a
browner tone, but several specimens of the present series closely approach
specimens of pyrropus from the Cameroon coast region.

Funisciurus congicus congicus (Kuhl)

Sciurus congicus Kun1, 1820, Beitr. Zodl., part 2, p. 66. Congo.

Represented by one specimen, adult male, Leopoldville, July 6, 1909.

Collectors’ measurements: Total length, 320 mm.; head ane body,
253; tail vertebre, 167; hind foot, 39.

Skull: Greatest Tenge 39.1; zygomatic breadth, 26.

This specimen is referable to the F’. congicus group, but whether it
represents the typical form is not now determinable. It is evidently not
F.. congicus interior Thomas, from Inkongo.

Tamiscus Thomas

Tamiscus Tuomas, 1918, Ann. Mag. Nat. Hist., (9) I, p. 38. Genotype, by
original designation, Sciwrus emint Stuhlmann.

Tamiscus (subgenus of Paraxerus) HouutstErR, 1919, U. 8S. Nat. Mus. Bull. 99,
part 2, May 16, p. 14.

Tamiscus emini emini (Stuhlmann)

Sciurus emini StuHLMANN (ex Matschie Ms.), 1894, ‘Mit Emin Pasha,’ I, part 1,
p. 320 (footnote), p. 321, fig. animal. NrumMANN, 1902, Sitzungsb. Ges. naturf. Fr.
Berlin, p. 180; ‘‘Liander zwischen Albert Edward und Albert Nyansa und nord-
westlich des Albert Nyansa bis Monbuttu.”’

1922] Allen, Congo Sciuride, Anomaluridx, Idiuride AY.

Parazxerus boehmi emini THomas, 1915, Ann. Mag. Nat. Hist., (8) XVI, Decem-
ber, p. 473. Medje (4), Mawambi (4), Poko (9 specimens).

Tamiscus emini emini Tuomas, 1918, Ann. Mag. Nat. Hist., (9) 1, January, p. 34.

Represented by 57 specimens (32 o”, 25 2, of which 8 are immature,
including 3 nurslings), from eleven localities, extending from Stanleyville
to Faradje, as follows:

Stanleyville, 10 (6 o&, 4 @, three of them immature), collected
August 11—28, 1909.

Batama, near Stanleyville, 1 (co adult), September 16, 1909.

Avakubi, 6 (3 #7, 3 2, all adult), October 8, 12, 13, 22, November 9,
1909, and January 22, 1914.

Ngayu, 3 (1 o, 2 Q, all adult), December 16-24, 1909.

Bafwabaka, 7 (4 co’, 3 @, all adult), December 28-31, 1909, and
January 5, 7, 1910.

Medje, 10 (6 &, 4 2, of which 4 are immature, including 2 nurs-
lings), January 15-20, 1910 (7, all adult but one), March 24, 1914 (a
nursling), and September 4, 9, 1914 (both immature, 1 a nursling).

Gamangui, 1 (Q adult), February 4, 1910.

Pawa, 1 (co adult), October 20, 1910.

Niangara, 5 (3 o’, 2 @, all adult), November 7—15, 1910, and May 18,
1913.

Faradje, 1 (o' adult), December 2, 1911.

Niapu, 12 (8 &, 4 9, all adult), November 14-25, 1913.

Collectors’ Measurements of Thirty-five Adults

No. of
Locality |Speci-| Total Length |Head and Body} Tail Vertebre | Hind Foot Ear
mens

Stanleyville| 7 |284(271-300)|127 (119-135) |/158 (150-170) |35 .0(84—36)|13 .0(12-14)
Avakubi 5 |277(269-287) |125 (122-128) |156 (147-165) |34 . 5(83-37)|13 .3(12-14)
Niapu 12 |276(267—292)|127(120-137) |148(1380-159)|32.6(31-35)|13 .6(12-15)
Medje 6 |282(275—283) |126(123-130) |154(152-158) |34.0(33-36)|13 .6(13-14)
Niangara 5 |275(252-294) |128(121-141) |149(141—-156) (33. 4(31-35)|13 .0(12-14)

Skull Measurements (Thirty-three of the Specimens in Table Above)

Locality No. of Specimens Greatest Length Zygomatic Breadth
Stanleyville 6 35.0 (84.3-35.9) 19.5 (19.1-19.7)
Avakubi 5 33.8 (33.1-34.8) 19.4 (19.1-19.7)
Niapu 12 34.4 (33.1-35.0) 19.1 (18.3-19.6)
Medje 5 34.2 (83.7-34.9) 19.3 (18.6-19.7)
Niangara 5 34.2 (32.9-34.9) 19.2 (18.5-19.8)

D8 Bulletin American Museum of Natural History [Vol. XLVII

The discrepancies in the average size at the different localities given
in the table is doubtless ascribable to differences in the average age of the
specimens.

This large series, from a wide range of localities, is astonish’ uni-
form in coloration, Stanleyville and Niangara specimens being indistin-
guishable. Season and age appear to exert little influence on coloration
of adults, excluding a few specimens in obviously worn pelage. In two
nurslings the general coloration of the upperparts is slightly more yellow-
ish than in adults and the black dorsal stripes are more sharply defined,
owing to the shortness of the pelage.

Tamiscus alexandri (Thomas and Wroughton)
Funisciurus alexandri THOMAS AND WrouGHTON, 1907, Ann. Mag. Nat. Hist.,
(7) XIX, May, p. 376. Type locality, Gudima, Iri River, Upper Uele (2specimens).
Paraxerus alexandri THomas, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 473. Medje (4), Poko (1 specimen).

Tamiscus alecandri THomas, 1918, Ann. Mag. Nat. Hist., (9) I, January, p. 37.

Represented by 19 specimens ( 8c’, 11 2), from 8 localities (Ava-
kubi to Faradje), collected as follows:

Avakubi, 4 (1 0’, 3 2, 3 adult, 1 9 embryo i in alcohol), November
23, 1913, and January 3, February 22, September 3, 1914.

Negayu, 2 (@, o, adult), December 22, 24, 1909.

Gamangui, 5 (1 o’', 4 @), January 28, February 8-11, 1910.

Medje, 2 (co, 9), April 1, May 25, 1914.

Pawa, 1 (co adult), October 10, 1910.

Nala, 1 (@ in alcohol), July 1913.

Rungu, 1 (2 adult), October 28, 1910.

Faradje, 3 (o’, all adult), November 29, December 2, 1911.

Collectors’ measurements of 4 adults (1 o’, 3 9) from Gamangui:
Total length, 215 (214-217); head and body, 105.5 (105-107); tail
vertebre, 110 (109-112); hind foot, 30 (29-31); ear, 14 (all 14).

Skulls of the same specimens and one other from same locality:
Greatest length, 29.5 (29.3-29 9); zygomatic breadth, 17.8 (17.3-18.3).

Collector’s measurements of 3 adult males from Faradje: Total
length, 219 (212-226); head and body, 107 (104-109); tail vertebre,
112.3 (108-117); hind foot, 29.7 (29-30); ear, 13 (12-14).

Skulls of the same specimens: Greatest length, 29.7 (29.5-29.9);
zygomatic breadth, 17.9 (17.4-18.6).

Collectors’ measurements of 9 specimens from other localities (Ava-
kubi, 3; Pawa, 1; Rungu, 1; Medje, 2; Ngayu, 2): Total length, 217
(210-230); head and body, 104 (96-107); tail vertebree, 112 (103-118);
hind foot, 29.9 (29-31); ear, 13.5 (12-15).

1922] Allen, Congo Sciuride, Anomaluridx, Idiuride 59

Skulls, 8 of the same specimens: Greatest length, 28.9 (28.1—29.4) ;
zygomatic breadth, 17.4 (16.9-17.7).

The middle of the back between the dark stripes is pale fulvous,
yellower than the sides of body, bordered on each side by a black and a
dull yellowish white stripe, and in many specimens indistinct traces of a
short blackish stripe outside of the whitish one. In the November,
December, January, and February specimens the black and white
stripes are sharply defined but in April, May, and October they are usu-
ally much less distinct owing to fading and wear. The white border of
the ears is at all times a conspicuous feature.

PRoToxERus Major
Protoxerus (subgenus of Xerus) Masor, 1893, Proc. Zo6l. Soc. London, (June 1),
p. 189, Pl. vin, figs. 7, 8, Pl. rx, figs. 7, 8. Genotype, by subsequent designation
(Thomas, 1897), Sciurus stangeri Waterhouse.

Protoxerus stangeri centricola (Thomas)

Sciurus stangert centricola Tuomas, 1906, Ann. Mag. Nat. Hist., (7) XVIII,
October, pp. 295, 297. Type locality, Katabi, Entebbe, Uganda (6 specimens).

Protoxerus stangeri centricola THoMAs, 1915, Ann. Mag. Nat. Hist., (8) XVI,
December, p. 473. Moera (3), Alimasi (2), Mawambi (2), ‘‘Poko’’! (13 specimens),
Belgian Congo.

Represented by 54 specimens (30 males, 24 females); all adult but 4;
among them 2 skeletons and 2 alcoholic; taken as follows:

Stanleyville, 1 (c”), September 30, 1914.

Kamunionge, southeast of Bafwasende, 1 (o”), September 21, 1909.

Avakubi, 5 (1 o%,2 9), June4, August 11, 26, 1914; (2 &, alcoholic),
August 20, 24, 1914.

Negayu, 7 (3 o&’', 4 9), December 11—238, 1909.

Bafwabaka, 2 (co, 2), December 30, 1909, and January 7, 1910.

Gamangul, 4 (3 o’, 1 2), January 29, and February 8, 9, 20, 1910.

Medje, 6 (2 o', 4 2), January 20, 23, March 15, September 10, 15,
1910, and February 27, 1914.

Niapu, 20 (12 o’, 82), November 14-30, December 2, 19, 1913,
and January 2, 1914.

Akenge, 5 (2 co’, 3 2), October 1-17, 1913.

Niangara, 2 (o’, 2), November 12, 19, 1910.

Faradje, 1 (o*), November 30, 1911.

1Mr. Lang believes that Dr. Christy’s specimens recorded as from ** Poko’’ were really taken in the
forest belt, farther south, toward Niapu.

60 Bulletin American Museum of Natural History [Vol. XLVII

As indicated above, 20 of the 54 specimens were taken at Niapu,
all but three during the last half of the month of November; all were
adults in fresh pelage. Niapu is about 60 miles south of Poko, from which
locality a large series collected by Dr. Christy has been referred by
Thomas (loc. cit., 1915, p. 473) to this subspecies.

Collectors’ measurements of the Niapu series (12 males, 8 females):

Total Length Head and Body Tail Vertebre Hind Foot Ear

o& 6587 (551-628) 284 (271-309) 301 (270-350) 69.8 (65-75) 22.3 (21-24)
Q@ 6591 (584-611) 283 (277-297) 307 (800-314) 69.8 (65-73) 22.0 (21-23)
Skulls, same specimens:
Greatest Length Zygomatic Breadth
( =occipito-nasal length) (=greatest breadth)
S& 67.23 (64.8-68.7) 37.5 (35.8-38.5)
Q 66.6 (64.6-68.7) 37.6 (36.3-38.1)

Collectors’ measurements of 15 adults (8 males, 7 females) from
other localities (Avakubi 3, Bafwabaka 1, Stanleyville 1, Gamangui 3,
Kamunionge 1, Medje 3, Ngayu 1, Niangara 2):

Total Length Head and Body Tail Vertebre Hind Foot Ear
3S 577 (557-590) 280 (261-296) 290 (277-314) 69.0 (68-71) 22.0 (21-24)
Q 586 (570-610) 290 (284-296) 292 (280-315) 68.4 (65-71) 21.7 (20-23)

Skulls, same specimens:

Greatest Length Zygomatic Breadth
(=occipito-nasal length) (=greatest breadth)
o 66.5 (64.6-67.0) 36.7 (35.5-37.8)
Q@ 66.2 (64.5-68.5) 36.8 (35.4-88.0)

The external measurements of the head, body, and tail! of the type
of centricola (an old female from Entebbe), ‘‘taken on the skin,” consid-
erably exceed the averages given above, but the foot, allowing 5 or 6 mm.
for the claws, is about equal to the smallest records of our specimens
measured in the field, and the greatest length of the skull (66.5 mm.)
differs less than a millimeter from the average of the 35 adult specimens
from the Belgian Congo given above.

The coloration of this large series is rather uniform, half-grown speci-
mens differing scarcely at all from the adults. A few of the latter, in
somewhat worn pelage, are a little pale from evident bleaching. The
extension of the gray of the dorsal region forward upon the head varies
somewhat, in some specimens gray-tipped hairs covering the crown as
far as the eyes, in others only as far as the front base of the ears.

1“ Head and body (overstretched) 310 mm.; tail, 330; hind foot, 61.’’

1922] Allen, Congo Sciuride, Anomaluride, Idiuridx 61

Protoxerus stangeri signatus Thomas

Protoxerus stangeri signatus THoMAS, 1910, Ann. Mag. Nat. Hist., (8) V, January,
p. 85. Type locality, Lodja, Upper Lukenie River, Belgian Congo.

A single specimen from Bolobo (skin without skull), presented to the
Expedition by Dr. Gerling, is apparently referable to this form which, as
indicated by the description based on the type specimen from Lodja, it
closely resembles. The type locality is some 400 miles east of Bolobo.
The differences from the series of P. s. centricola as recorded above are

‘slight and it is here recognized mainly on geographical grounds.

Euxerus Thomas

‘Euxerus THomas, 1909, Ann. Mag. Nat. Hist., (8) III, June, p. 473. Genotype,
by original designation, Sciurus erythopus Geoffroy.

Euxerus erythopus lacustris (Thomas)

Xerus erythropus lacustris THomas, 1905, Ann. Mag. Nat. Hist., (7) XV, April, 3
p. 388. Masindi, Unyoro.

Eucxerus erythropus lacustris THomas, 1915, Ann. Mag. Nat. Hist., (8) XVI,
December, p. 474. ““Panga”’ (near Poko) (3), Poko (8 specimens).

Represented by 30 specimens, collected as follows:

Faradje, 20 (2 ¢ and 10 @ adults, 1 nursling, 7 one-fourth to one-
third grown), February 20, 22, 26, March 4, 5, 14, 31, April 3, 11, 29, 30,
May 27, June 29, September 3, 11, 1911, October 16, December 14,
1912, January 10, 12, 1913.

Niangara, 9 (2 adult, 7 immature, of which 5 are nurslings), Nov-
ember 12-28, December 22, 1910, and January 2, 1911.

Rungu, 1 (* adult), January 30, 1913.

Collectors’ measurements of 11 adults (2 males, 9 females) from

’ Faradje: Total length, 492 (474-515); head and body, 282 (259-297) ;

tail vertebree, 209 (189-230); hind foot, 72 (68-75); ear, 18.8 (18-20).

Skulls, same specimens: Greatest length, 65.1 (63.6-67.2); zygo-
matic breadth, 34.2 (33.7-34.6).

The three localities at which specimens were taken are all in the
open districts of the savannah in the northeastern Belgian Congo.

Young specimens a few weeks old are similar in coloration to the
adults, the pattern being the same, but a little lighter in tone, the light
tips to the hairs of the upperparts, owing to the shortness of the pelage,
concealing the darker basal portion. The tail, however, is externally
white, the long white tips of the hairs usually wholly concealing the broad
subapical black zone of the tail hairs. Later, as the animal increases in
size, the black base of the hairs forms a narrow black median line on both

62 Bulletin American Museum of Natural History [Vol. XLVIT

the upper and the lower surfaces of the tail; still later, in specimens one-
third to half grown, both surfaces of the tail are grizzled black and white,
with the sides and tip white, and the body pelage, in texture and colora-
tion, has become like that of adults, the juvenal coat having been re-
placed by molt.

ANOMALURIDZ

The one hundred and twenty-five specimens of Anomaluridz repre-
sent three of the four superspecific groups of this family proposed by
Matschie in 1914,! and are referable to three forms.

ANOMALURUS Waterhouse

Anomalurus WATERHOUSE, 1842, Ann. Mag. Nat. Hist., X, pp. 201, 202; 1842,
Proc. Zoél. Soc. London, (January 1843), pp. 124-127. Genotype, by monotypy
(also by original designation), Anomalurus fraseri Waterhouse.

Aroxthrus WATERHOUSE, 1842, Proc. Zod]. Soc. London, (January 1843),
p. 124, footnote. Substitute name to replace Anomalurus Waterhouse in case the
latter is found to be preoccupied.

Anomalurus, as restricted by Matschie (loc. cit., 1914), includes about a dozen
forms, the greater part of which are subspecies of A. frasert.

Anomalurus jacksoni jacksoni de Winton

Anomalurus jacksoni DE WINTON, 1898, Ann. Mag. Nat. Hist., (7) I, March, p-
251. Ntebe (=Entebbe), Uganda.

Anomalurus jacksoni THoMAs, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 472. Moera (1), Medje (2), Poko (5 specimens).

Represented by 58 specimens (25 males, 29 females, all adult but 3,
including 4 in alcohol, 4 skeletons, and a skull), collected as follows:

Avakubi, 2 (co, @ in alcohol), March 31, April 14, 1914.

Medje, 28 (12 &%, 18 2; 2 in alcohol, 4 skeletons), January 20-24,
March 6-21, April 9-26, May 2, August 1, 3, September 9, 16, October -
6, 1910.

Gamangui, 1 (skull only), February 1, 1910.

Niapu, 16 (10 &, 6 2), November 12-18, December 2-19, 1913.

Akenge, 7 (o’', 6 9—1 @, embryo in alcohol), September 29-30,
October 9, 11, 28, 31, 1913.

Panga, 4 (o’, 3 2 ), September 14-18, 1914.

Collectors’ measurements of 16 adult specimens (9 co’, 72) from
Medje:

Total Length Head and Body Tail Vertebre Hind Foot Ear

& 549 (518-563) 317 (296-332) 2387 (228-250) 61 (58-63) 38.5 (386-41)
Q 582 (559-621) 331 (319-342) 258 (240-280) 63 (62-65) 40.0 (389-41)

“Bin neuer Anomalurus von der Elfenbeinkiiste.’ Von Paul Matschie. 1914, Sitzungsb. Ges.
naturf. Freunde Berlin, No. 7, July, pp. 349-351. (1) Anomalurus Waterhouse, (2) Anomalurodon,
(3) Anomalurops, (4) Anomalurella.

1922] Allen, Congo Sciuride, Anomaluride, Idiuridz 63

Skulls, same series:

Occipito-nasal Length Zygomatic Breadth
Gui50-0) (ba15-54.3) ; 36.9 (35.0-38.0)
OP ai lepop-OSeo) 38.3 (35.0-39.8)
Collectors’ measurements of 15 adult specimens (8 o’, 7 2) from
Niapu: 7
Total Length Head and Body Tail Vertebrz Hind Foot Ear

& 559 (540-570) 313 (298-323) 248 (222-267) 60.4 (58-65) 37.6 (35-39)
Q 582 (554-628) 311 (301-344) 261 (252-284) 61.3 (62-64) 38.1 (36-40)

Skulls, same series:

Occipito-nasal Length Zygomatic Breadth
6100.0) (One 2-017 28) 37.5 (36. 1-38 .0)
0757-2 (56,0-58 .9) 37.5 (36.8-38.4)

The three adults (1 &, 2 9) from Panga agree in proportions and
measurements with the Medje and Niapu specimens. The single speci-
men from Avakubi is a young adult male.

In compiling the measurements given above only specimens in which
the dentition was fully mature were used; but while the relative number
with unworn teeth varies in the different categories, and tends to lower
the average for the series when they predominate, this factor, in the
present series, does not account for the slightly larger average size of the
females in the above statistical summaries. In the discarded specimens
the last molar was not fully developed, varying in different specimens
from just breaking the alveolus to one-half to two-thirds full height, but
still unpigmented. In such specimens the total length of the skull
ranges from 50 to 53 mm., as against 55 to 58 mm. in adults.

ANOMALURELLA Matschie

Anomalurella Matscuim, 1914, Sitzungsb. Ges. naturf. Freunde Berlin, July,
p. 351. Genotype, by original designation, Anomalurus pusillus Thomas.

Anomalurella pusilla (Thomas)
Anomalurus pusillus THomas, 1887, Ann. Mag. Nat. Hist., (5) XX, December,
p. 440; 1888, Proc. Zool. Soc. London, p. 8, Pl. 1, animal. Bellima, 1 @ (type);
Tingasi, 1 o.
Anomalurus pusillus THomas, 1915, Ann. Mag. Nat. Hist., (8) XVI, December,
p. 472. Moera (1), Medje (4), Poko (7 specimens).

Represented by 53 specimens (21 o’, 29 2), of which 5 are immature
(but none very young), 2 are skulls only, 4 skeletons, and 2 in alcohol,
collected as follows:

Avakubi, 1 (2), September 18, 1913.

Ngayu, 2 (2 2), December 17, 20, 1909.

64 Bulletin American Museum of Natural History [Vol. XLVII

Medje, 36 (12 #7, 21 2—2 &, 2 2 immature, 2 skulls, 4skeletons, 1
embryo in alcohol), January 16-26, March 8-26, April 4-27, May 14,
_ June 30, August 18, October 7, 1910, March 23, June 25, 1914.

Niapu, 10 (7 &, 3 2), November 19, 24, 26, December 4-9, 16,
1913. |

Akenge, 4 (2 o, 2 9), September 30, October 14, 16, 1913.

Collectors’ measurements of 23 adults (7 o', 16 2) from Medje:

Total Length Head and Body Tail Vertebre Hind Foot Ear
o 371 (359-390) 227 (210-242) 144 (138-152) 43.0 (40-46) 29.8 (30-32)

Q 373 (357-401) 230 (212-246) 148 (141-157) 44.5 (40-47) 30.0 (30-32)
Skulls (10 o&, 14 2), same series:
Occipito-nasal Length Zygomatic Breadth
co 44.8 (48.6-46.6) 29.8 (28.0-30.9)
Q 45.2 (43.6-47.6) 29.9 (28.0-30.8)

Collectors’ measurements of 9 adults (7 o’, 2 9) from Niapu:
Total length, 371 (853-382); head and body, 219 (213-227); tail verte-
bre, 143 (130-147); hind foot, 43.7 (42-46); ear, 29.2 (28-30).

Skulls, same specimens: Occipito-nasal length, 45.1 (43.5-46.3);
zygomatic breadth, 29.7 (28.3-30.2).

Very few of the specimens of the present series conform very closely
in the coloration of the upperparts to the original description and accom-
panying colored plate of the species, of which it issaid: ‘‘General colour
above uniform dark grizzled gray, the tips of the hairs forming a terminal
band of pale gray or olivaceous.’”’ In many specimens of the present
series this ‘‘terminal band”’ is near ochraceous rufous, intensified in ex-
ceptional specimens to pale tawny. One specimen from Akenge may be
noted as having the pelage of the back hazel for the terminal haif or more
of the hairs, recalling forcibly the dorsal region in A. beldeni Du Chaillu
(=? erythronotus Milne-Edwards) but the red is browner. This speci-
men, however, is probably abnormal, as the new coat, coming in in
patches, conforms to that of normal specimens.

ANoMALUROPS Matschie

Anomalurops Matscutn, 1914, Sitzungsb. Ges. naturf. Freunde Berlin, No. 7,
July, p. 351. Genotype, by original designation, Anomalurus beecrofti Fraser.

This group includes six described forms, the greater part of which
appear to be subspecies of A. beecrofti. The type locality of A. beecroftr
chapint, described below, is far to the eastward of any previously known
locality for the beecrofti group.

1922] Allen, Congo Sciuride, Anomaluride, Idiuride 65

Anomalurops beecrofti chapini, new subspecies

Type, No. 50480, & adult, Medje, Belgian Congo, May 6, 1910; Herbert Lang
and James P. Chapin. American Museum Congo Expedition. Orig. No. 861. Named
for James P. Chapin.

Smaller and much lighter in color than A. beecrofti citrinus Thomas,! from Benito
River, Spanish Guinea.

General color above (including membranes), yellowish gray; middle of back
from occiput to sacral region with an irregular broad band of ochraceous orange,
varied with black, the hairs individually (about 20 mm. in length) mouse-gray
basally, passing gradually into a broad band of dull black, followed by a subapical
band (5-6 mm. wide) of ochraceous orange and conspicuously tipped with black;
hairs of upper surface of membrane for the antero-lateral fourth of the border rigid
and intense black; a well-defined patch of white or buffy white at lateral base of
ears, indistinctly merging by a downward sweep with the white or whitish half collar
in front of shoulders, and the usual small tuft of partly concealed white hairs on occi-
put; front and sides of head dull gray or buffy gray, extending laterally to sides of
throat; general color below (including membranes) ochraceous buff to base of hairs
(varying in some specimens to light buff or even whitish), except on the throat and a
broad median band thence to anal region (usually narrowing posteriorly) and the
inside of thighs, which parts are intense orange-rufous in high-colored specimens,
paler in others; basal third of tail and caudal membrane below like ventral surface,
above like the lower back; apical two-thirds of tail dull brown, varying in different
specimens from dark brown to pale fulvous brown and even cinnamon-brown;
upper surface of feet dull grayish with a slight buffy suffusion; soles and palms naked,
pale brown, as are the nose and apical two-thirds of ears.

Collectors’ measurements of type: Total length, 512 mm.; head ent body, 310;
tail vertebre, 202; hind foot, 59; ear, 29.

Miisurerments of skull (type): Greatest (occipito-nasal) length, 54.5; condy-
loincisive length, 51.4; zygomatic breadth, 34.4; least interorbital breadth, 18.4;
breadth of brain-case, 26.3; length of upper toothrow, 11.9.

Collectors’ measurements of 10 adults (5 o&, 5 9), of which 8 are from Medje
and 1 each from Poko and Akenge:

Total Length Headand Body Tail Vertebre Hind Foot Kar
o 512 (501-518) 301 (278-323) 211 (195-223) 59 (56-62) 31 (29-32)
Q 510 (475-555) 297 (277-830) 213 (198-226) 59 (58-60) 31 (29-32)

Represented by 14 specimens (8 o, 6 9, all adult but 2), collected as follows:

Medje, 12, (6 &%, 6 9), March 13, April 9, 13, 27, 30, May 6, 8, September 24,
October 12, 1910, February 28, 1914.

Poko, 1 (o’), August 22, 1913.

Akenge, 1 (o), October 1, 1913.

Anomalurops beecrofti chapini is geographically nearest A. bee-
croft. citrinus Thomas (type from “Benito River, Spanish Guinea,”’
collected by G. L. Bates), from which it differs in considerably smaller
size and much less intense coloration. Fortunately I am able to make

tAnomalurus beecrofti citrinus Thomas, 1916, Ann. Mug. Nat. Hist., (8) XVIII, August, p. 236.
‘About a dozen specimens examined.

66 Bulletin American Museum of Natural History [Vol. XLVIIL

direct comparison of the Congo series with four topotypes! of cztrinus
also collected by Mr. Bates. As shown in the accompanying tabulated
measurements, A. 0. citrinus exceeds A. b. chapini in total length by
about 50 mm., and about the same in head and body length, while there
is very little difference in the length of the tail. (It is probable that these
latter measurements were not taken by the same method in the
two cases.) In skull length citrinus exceeds chapini by about 4 mm.,
with the other skull measurements proportionately different. The
‘mpression given by comparison of the two series is a greater size differ-
ence than the measurements indicate, the citrinus skulls being more
heavily ossified, with stronger ridges for muscular attachment in skulls
of equal age than is the case in chapini. The color differences are strongly
marked. The ‘ochraceous olive” or ‘‘citrine drab” effect above in
citrinus is replaced by light clear gray, or slightly yellowish gray, in
chapini, with a corresponding difference in the tone of the median dorsal
band; below the ferruginous tone is much darker in the former, approach-
ing chestnut-red on the throat and median line in cérinus in place of
orange-rufous in chapini.

The series of 14 specimens of A. b. chapini presents the usual wide
range of individual variation in both size and coloration. The smallest
specimen in cranial measurements is a female (skull, 51.8 X 32.9) in
which all the cheek-teeth have attained full development but are un-
worn; in external measurements it is the largest of the females except
one, which is much the largest specimen of the entire series (skull, 58.5 <
36.8) in both external and cranial measurements.

The color above varies from clear light gray to yellowish gray, and
the rufous dorsal line is in some weakly developed or nearly obsolete,
in others heavy and continuous from the crown to the hips. The broad
rufous zone of the median underparts likewise varies greatly in extent
and intensity—from orange-rufous to dark ferruginous, and the adjoin-
ing lateral parts from ochraceous orange to pale buff. Young specimens
are much paler below than the adults. The white crown spot is nearly
always plainly distinguishable and usually forms a distinct mark which is
occasionally conspicuous. In one specimen it is a transversely-oval
patch, 15 X 25mm. in area. There is apparently no sexual difference in
size or color.

1Borrowed from the United States National Museum through the kindness of Mr. G.S. Miller, Jr.,
Curator of Mammals.

1922] Allen, Congo Sciuride, Anomaluride, Idiuride 67

Collectors’ Measurements

Locality Sex Total and Verte- | Hind Foot
Length! Body | bre

‘Benito River,

Type! A. b. citrinus ‘Spanish Guinea| @ HSIe SOOT ea 220 ay OO SU:

84546 N. M. A. b. citrinus| e: Q Hop a wooWuies clo) ar ss

84547 “ i ee ot rt of FAO" 35be | S54, 53 <

84548 “ ci * ob Q 585 | 380 | 205 | 56 a

Average, 4 specimens 568 | 361; 208 | 56.5 “
Medje,

50477 A. b. chapini ‘Upper Congo Q 515 | 305 ALO] 3S) CoUs
50480 “ se ca os of SIA SlOh 205" 59 :
50481 “ i: ie ay Q 5551. 3380 | 225 | 60 a
50482 “ te - of BIZ |. 298 | 249 | 62 Z|
50483“ ee e Q 490 | 282 | 208 | 60 a
mee fe e 2s of 518 | 323 195 | 60 os
50485 “ ie ‘Poko of HO 208 siete o9 aoe

Average, 7 specimens | 517 | 305 | 211 | 60 a

Measurements of Skulls

os Sex | Length | Aiect | recthcow
Benito River,
Type! A. b. citrinus Spanish Guinea 2 58.9 38.0 12.8
84546 N. M. A. b. citrinus a ie Q 59.5 38.3 13.0
845475,“ Ke fs re £ of 60.5 | 38.4 11332
84548 =“ S i - a Q 59.7 38.3 t3e0
BAe Y es 2 oh Q 58 .0 Ba Al il tnsez
Average, 5 specimens 59.2 38.1 13.0
Medje,
50477 A. b. chapini Upper Congo Q SH) 2 35.8 12.6
SO4R0). *% e = A of 54.3 33.8 2?
HOASIy = i es i Q 58.5 36.8 WPF
50482 “ as be of ot 56.4 36.2 12.5
50483“ eS i i" Q 52.6 355 08) 1250
50610 “ o e 9 of 55.0 36.3 117
50485“ Me Poko of 55.1 33.8 11.8
Average, 7 specimens 55.3 35.4 12.2

1From the author’s description (loc, cit.).
2Toothrow abnormally short and omitted from the average.

68 Bulletin American Museum of Natural History [Vol. XLVII

IDIURIDE

The Idiuride, recently separated from the Anomaluride as a
distinct family group by Miller and Gidley, are represented by three
quite different forms, two of which are here for the first time described.

Ip1urus Matschie

Idiurus Matscuig, 1894, Sitzungsb. Ges. naturf. Freunde Berlin, No. 8, August,
pp. 194-200, 1 text-fig. Genotype, by monotypy, Idiurus zenkeri Matschie.

Idiurus zenkeri zenkeri Matschie

Idiurus zenkeri Matscuin, 1894, Sitzungsb. Ges. naturf. Freunde Berlin, No.
8, October 16, p. 197, text fig. p. 198 (animal). Type locality, Yaunde Station,
Cameroon District, West Africa. One specimen.

Represented by 30 specimens (22 skins with skulls, 8 in alcohol),
collected as follows:

Medje, 27 (14 o&, 13 2; 21 skins and skulls, 6 in alcohol), January
21, 25, March 9, 16, 1910.

Avakubi, 1 (@, skin and skull), January 22, 1914.

Niapu, 2 (1 &, 1 2, in alcohol), January 27, 1914.

Collectors’ measurements of 19 adults (10 males, 9 females) from
Medyje:

Total Length Headand Body ‘Tail Vertebre Hind Foot Ear
or 170 (165-175) 71 (64-78) 99 (93-104) 17.0 (16-18) 13.6 (12-14)
Q 173 (160-187) 73 (65-86) 101 (95-108) 17.3 (16-18) 138.4 (12-14)

Measurements of 14 skulls (7 males, 7 females), from the same
series:

Greatest (=occipito-nasal) Length Zygomatic Breadth
ot 21-0: (20. 5-213) UPA A AA MLZ)
Oe Ae (Zea lke 8) 12:4 (G1 79-1278)

This fine series, particularly the 21 skins from Medje, throws much
light upon questions of individual, sexual, and seasonal variation. The
measurements, both external and cranial, indicate a slightly larger
average size for females than males, but there is no recognizable sexual
difference in coloration.

The range of color variation is considerable in the Medje specimens
taken at the same date, due largely to the condition of the pelage in
respect to wear, the general tone of the coloration becoming darker as
the tips of the hairs wear off, showing more of the basal fur, while the
hair-tips become paler by fading. Comparison of the twelve specimens

1‘Synopsis of the supergenerie groups of Rodents.’ Gerrit S. Miller, Jr., and James W. Gidley,
1918, Journ. Washington Acad. Sci., VIII, No. 13, July 19, p. 422.

1922] Allen, Congo Sciuride, Anomaluridx, Idiuridz 69

taken March 16 with the eight taken January 25, shows that the latter
average darker in general effect and the hair-tips paler, yet certain speci-
mens of the January series can be matched exactly by the paler speci-
mens of the March series. The hair-tips on the back of the brighter
colored examples of the March series are near snuff-brown, varying in
intensity in different individuals, and about cinnamon-buff on the ventral
surface, but often nearly wanting through wear, as in the single Avakubi
specimen taken January 22, the most worn of any of the entire series of
twenty skins.

A single skin and skull! of I. zenkeri, from the southern Cameroon,
and thus practically a topotype, is rather darker than the average of the
Medje series, but differs so little from some of them that they are provi-
sionally referred to this species. Their relationship to I. zenkeri kivuenszis,
recently described by Lonnberg,? is not at present determinable. It
appears to be a much darker form than typical zenkerz.

Idiurus langi, new species .
Plate V

Type, No. 50542, & adult, Medje, Belgian Congo, March 16, 1910; Herbert
Lang and James P. Chapin, American Museum Congo Expedition. Orig. No. 737.
Named for Herbert Lang, leader of the American Museum Congo Expedition.

Size of and proportions nearly as in Idiurus macrotis Miller, but very different in
coloration.

Upperparts (type, in fresh, unworn pelage) washed with clay-color (Ridgway,
1912), strongest on middle of back, less heavily on lower back and sides ; in worn
pelage much paler (about cinnamon-buff), the light hair-tips partly worn off (almost
wholly on lower back and sides); pelage of middle of back (in fresh coat), 11.5 mm.
in length, the buffy tips about 2 mm. long, followed by a dark zone of about equal
width, the basal two-thirds ‘‘mouse-gray.’’ Underparts heavily washed with warm buff,
almost wholly concealing the light neutral gray of the basal fur (varying in different
specimens, especially when worn) to a faint wash of lighter tone); a conspicuous pale
yellowish white patch on sides of nose extending from base of rictal bristles to the
naked nose pad, about 4 X 6 mm. in extent; chin and interramal region white or
pale yellowish white; upper surface of membranes thinly clothed with brownish black
hairs, under side nearly naked; membranes and ears pale brownish (ears in some
specimens slightly darker brown); feet and greater part of tail yellowish brown, the
long tail hairs dark brown with a faint tone of chestnut, much less dark than in J.
macrotis. Thescale pad on the ventral base of the tail is much longer than in macrotis
(given as 17 mm.), varying from 20 to 25 mm. in length, and the scales are larger and
tend to form regular rows, and beyond what may be considered as the “‘scale pad’”’
proper, the annulations on the lower surface of the tail are conspicuous and roughened,
so that in some specimens it is difficult to determine what should be regarded as the

1No. 125438, U.S. Nat. Mus., 2, Efulen, Bulu Country, Cameroon, July 21, 1903, coll.G. L. Bates.
21917, Kungl. Svens. ventensk. Akad. Handl., LVIII, No. 2, September, p. 67. Masisi, Belgian
Congo, about forty miles northwest of Lake Kivu. Twospecimens, adult and young.

70 Bulletin American Museum of Natural History [Vol. XLVII

apical end of the pad, as distinguished from the annulations. The fringes on outer
edge of both fore and hind feet, the small tufts of whitish bristly hairs at tarsal and
metatarsal joints, the tail fringes, and the scattered long hairs in the dorsal pelage,
are evidently generic characters, being common to the three forms of Idiurus here
under consideration.

Collectors’ measurements of the type!: Total length, 224 mm.; head and body,
94; tail vertebrx, 130; hind foot, 20; ear, 18.

Collectors’ measurements of type and 4 topotypes (all adult males): Total
length, 218 (207-224); head and body, 91 (86-94); tail vertebre, 129 (124-133);
hind foot, 21 (20-22); ear, 15.7 (14-18).

. Skull (measurements of type!): Greatest length, 26; zygomatic breadth, 16;
least breadth of frontals,? 6.6; greatest breadth of nasals,? 3.2; upper toothrow, 3.5;
distance between inner bases of m’, 1.4; do. m!, 1.1; greatest length of mandible,
16.4; greatest depth (at coronoid), 10.2; lower toothrow, 4.

Skull (type and same 4 topotypes): Greatest length, 25.8 (25.1-26.2); zygomatic
breadth, 15.5 (15.0-16.0).

Represented by 6 adult males, all of which are skins with skulls, and 1 adult female
in alcohol, all taken at Medje, January 25 (5 specimens) and March 16 (2 specimens),
1910.

The type is the only specimen in fresh, wholly unworn pelage. All the others
show more or less wear, especially on the lower back and sides, and they vary much
in the amount of buffy wash, both above and below, and form a graduated series from
clay-color to a pale tone of buff on the upperparts, and on the lower parts froma
strong yellowish wash te only a faint pale tone where the hair-tips are least worn.
The two March 16 specimens differ greatly from each other in coloration and amount
of wear; the January series of skins (all taken January 25) differs similarly in respect
to amount of wear and consequent tones of color on both upper and lower surfaces.

Idiurus langi is smaller than I. macrotis in external measurements,
but the cranial measurements are practically the same. It differs, how-
ever, strikingly in coloration, both above and below, the general color
being much lighter, especially in respect to the basal fur, ears, and mem-
branes. J. langi differs from I. panga in much larger size and in colora-
tion, especially of the ventral surface, which has a pinkish tone in panga
instead of yellowish, and the upper surface is much more heavily washed
with buff. The yellowish white, sharply defined nose spot of lang: will
alone readily distinguish it at a glance from either macrotis or panga.

Idiurus panga, new species

Type, No. 50605, @ adult, Panga, Belgian Congo, September 18, 1914; Herbert
Lang and James P. Chapin. American Museum Congo Expedition. Orig. No. 2552.

Similar to Idiwrus macrotis Miller’, but much smaller and considerably paler
throughout, including the basal fur.

1For measurements of I. macrotis see p. 71, where they are given in comparison with I. panga.
2Fronto-nasal sutures solidly ankylosed and indistinguishable, so that length of frontals and nasals

cannot be given.
3Tdiurus macrotis MiLLER, 1898, Proc. Biol. Soc. Washington, XII, pp. 73-76, figs. 15-19 (skull,

ear, foot, and tail). Efulen, Cameroon District, West Africa.

1922] Allen, Congo Sciuride, Anomaluridz, Idiuridx 71

Upperparts (in comparison with a para-topotype of J. macrotis) with the hairs
narrowly tipped with light drab (instead of “‘sepia’’), forming a slight wash of this
tone, strongest on middle of back and sides of neck, darkened by the deep neutral
gray (instead of dark plumbeous) underfur, which color predominates over the lower
back and flanks; underparts superficially pale vinaceous buff (instead of yellowish
wood-brown), the basal fur dark gull-gray (instead of plumbeous); upper surface of
membranes thinly clothed with dusky brown hairs (less dark than in macrotis, as
are also the membranes themselves) ; ears, feet, and base of tail also much lighter than
in macrotis.

Collectors’ measurements of type: Total length, 209 mm.; head and body, 73;
tail vertebre, 126; hind foot, 20 (20.5 in dry skin); ear, 18.

Collectors’ measurements of the type and 3 topotypes (1 male, 3 females): Total
length, 206 (199-212); head and body, 80.5 (73-87); tail vertebra, 123 (117-128);
hind foot, 20.5 (18-21); ear, 17.3 (17-18).

Corresponding measurements of the type and topotype (2 males) of macrotis,
as given by the author (loc. cit): Total length, type 241, topotype, 228; head and
body, 108, 105; tail vertebra, 133, 123; hind foot, 21, 22; ear, 16, 15.5.

Skull (measurements of type, with measurements of type and topotype of
macrotis in parentheses): Greatest length, 25 (26, 27); zygomatic breadth, 14.9
(15, 16); length of frontals,— (96, 98); least width of frontals, 6.5 (7, 7); length of
nasals,—! (7, 7); greatest breadth of nasals, 3.1 (3.25, 3.3); upper toothrow, 3.5
(3.8, 4); greatest distance between molars at m', 1.5 (2.2, 2); least distance between
molars at m!, 1 (1.2, 1.2); greatest length of mandible, 14.9 (15, 16); greatest depth
(at coronoid), 9.7 (10, 10.6); lower toothrow, 3.7 (4, 4).

Skull (type and 3 topotypes): Greatest length, 25.0 (24.-25.6); zygomatic
breadth, 14.5 (14.2-14.9).

Represented by 4 specimens (1 0, 3 Q), all from Panga and all collected the
same day, September 18, 1914. All are old adults, with the dentition fully mature
and the fronto-nasal sutures fully ankylosed and wholly indistinguishable.

Idiurus panga is a member of the J. macrotis group and is so
different from the J. zenkeri group as to need no comparison with it.
It differs from macrotis in smaller general size, much less heavy skull and
correspondingly weaker dentition. Also in the much paler hair-tips
above, the decidedly pinkish tone of the underparts, and the lighter
color of ears, membranes and feet, and also of the basal underfur, both
above and below. The ears appear to be decidedly larger than in macrotis,
as indicated by the field measurements and by direct comparison with
the para-topotype loaned me for examination through the kindness of
the describer of the species.

1Fronto-nasal sutures fully ankylosed and indeterminable.

Bens
» Fs,

(fel

Sa CaS Seen os ~ OrnrrHoLoey < us se .
Th Castes tien of ike: Weaver Birds. ‘By James P. Chapin, 1 1917,
oe ~ Bulletin, XXXVI, Art.9, pp. 248-280, Pls. vi-x, 10 text figures. —

: tin, XXXVI, Art. 12, pp. 327-334, |
a _Fidd. Notes on M. olothrus bonariensis and M. badins. ‘By Leo E. Miller,
ia 85 Ae Bulletin, XXXVIL, Art. 22, pp. 579-592. *

| = Seas

A ‘study. of ae Adlantic Oceanites. ‘By Robert Giditian ae 1918, e : as
ee 60c.
= “The Geographical Distribution of Color and of Other Variable Characters =

Bulletin, XXXVUEI, Art. 4, pp. 117-146, Pls. ran, 7 text figures.

in the Genus Junco: -a-New Aspect of Specific and Subspecific Values.

ooe=baa.

By Frank M. Chapman, 1921, Novitates, No. 2, pp. 1-8.

~ 1921, Novitates, No. 7, pp. 1-9.

ae Revision of Atlapetes guituralis with Deocipdons of Three N ew inc. By

Jonathan Dwight and Ludlow Griscom, 1921, ‘Novitates, No. 16, pp. 1-4.

me | | Notes: on a New Ox-pecker and Other Little-known Birds of the Congo. By : et

~ James. Pp. Chapin, 1921, Novitates, No. 17, pp. 1-16.

Brazil. By Frank M, Chapman, 1921, Novitates, No. 18, pp. 1-12,

Description of Proposed New Birds from Central America, with Noteson
— Other Little-known Forms. By Waldron DeWitt Bey ane oe ue

~ Griscom, 1921, N ovitates, No. 25, pp. 1-13. :
A Review of the Diving Petrels, Contributions fein the roger Santond

Collection. By Robert Cushman Murphy and Francis Harper, 1921, ae

~ Bulletin, XLIV, Art. 17, pp. 495-554, Pls. XX-XxIv, 7 text figures.

_ Descriptions BE Koons New Birds Sone Bolivia, Brasil, and | Venezuela.

"Descriptions of Proposed New Birds from Colombia, Ecuador, ‘Peru, and Se

_ Descriptions of New Birds from Santo. Domingo and Remarks on Othersin® =
the Brewster-Sanford Collection. ae Frank M. Chapman, EG Bulle- ae

38005

* : 7 fe be S
ee A New Albatross from the West Coast of South America. By Rohe Cush 2s
— “man Murphy, 1917, Sellen, eke Att. 35, Pee 861-864, I text

~ By Jorathan Dwight, M.D., 1918, bec, XXXVI, At. 9; oP. 269—-

--309; Pls. xi-xi, 5 maps.. ee
_ Desiptions of Proposed ‘New Birds: from Peru, Bolivia, Accontnn. and i

Chile. - By Frank. M. Chapman, 1919, Pes XL, Art. 5 pp. =

eee

_ Descriptions of Proposed New Birds from Brazil, Paraguay, and ieecitnn ee

if ; “By George K; ‘Cherrie and (Mrs.) Elsie M. B. Beschoaheties, 1921, ©

-Novitates, No. 27, pp. 1-6.

The Birds of The American Mosaun of N sea History: 8 Asiatic. yaslnclak
~ Expedition of 1916-1917. _ By Outram Peale. es Bulletin, = V54

Art. XX, pp; 575-612.

~M. Chapman, 1922, Novitates, No. 30, pp. 1-15.

Zs -Deseriotions of Apparently New Birds from Cclombin Ecuador, ad

_ Argentina. By Frank M. Chapman, 1422, Novitates, No. 31, pp. 1-8.

“The Species and Geographic Races of Steganura. - BY James P: hee 5

1922, Novitates, No. 43, pp.1- 12

Cae of a. New Race of the Lesser Blick backed: Gull, from the aS

_ Azores. oy Jonathan pes 1922, No, 44, ‘Pp. 4,2.

The Distribution of the Seatiows of the Genus Pygochelidon. By Bronk a

sS pies nae | Mmmorms

oe ses : Vohins 2 Spelt and Paleontology.
noe Volumes U-VII. Anthropology. *. eae
Sheets ue ae Volume IX.” ‘Zoology and Paleontology. -
Bi Pos _ Volumes X-XIV. Anthropology. — os
Be Il, IV, V, VII, VII, x =XIV, and an Ethnographical a
Volumes: I-x of the Memoirs of the oe North Pacific breiecaer

Tei

- Memoms—New Sunms

yl I and II. Zoology. and Paleontology.
Volume Hl, part 1. Entomology. ou
Volume Hy, ied 1-2 Be oe

Bowen see eo rae

| parte 1-4; 2x parts 1-3; XXIV, pars 1-8; XXV, pat 1; a 1
‘Monocraras. tae oe = ae a

A Review of the Primates. ‘By D. G. Elliot. 38 volumes.

) By E.. D. . Cope and W. D. Matthew.

NaTouRAL , History
- Journal of The American Museum of N atural I History

Museum of Natural Hickory issued bimonthly.
HANDBOOKS. Numbers 1-8. i os soe Ss a — ia oe
GuipEe LEAFLETs. Numbers 1-54. oe

ANNUAL Ruronrs, ‘First (1869) +0 Pity hid oan.

tion to ihe omtaiees of the Museum,

My Moog Boat
- “THe AMERIC. \N MUSEUM
a ‘e conco EXPEDITIC N MANATEE AND
ce _ OTHER peed MANATEES
©

POS ROBERT T. HATT

BULLETIN
OF
‘THE AMERICAN MUSEUM OF
NATURAL HISTORY © |

VOLUME LXVI, 1934,
ARTICLE IV

NEW YORK
September 10, 1924

59.9, 55 M (67.5)
Article IV.—A MANATEE COLLECTED BY THE AMERICAN
MUSEUM CONGO EXPEDITION, WITH OBSERVATIONS
ON THE RECENT MANATEES!

By Rosert T. Harr

PuatE XXVII; Text Ficures 1 anp 2

CONTENTS

PAGE
Mier yD) GO Nee eee ne ee ee ee Gee ee aria MN See Le 033
INOee NCHARURE On THE VIAN ATHENS: 00. sock soils own scala douse ee wee nie ees 034
CRITERIA FOR THE CLASSIFICATION OF THE MANATEES....................... 539
THE DISTRIBUTION OF T'richechus IN THE AFRICAN REGION................... 554
UNCERTAIN OR ERRONEOUS Reports oF 7’. senegalensis................ 588
‘STIR AUEYZ. oo ere ce OR bc. tne DM fe SUR oP A el ge 560
LETUBTE OGTR NETS 2079 sR ae SRR er MER CP 562
ADDENDA REGARDING LAKE CHAap MANATEES ...... Ree Pek Seer OG

INTRODUCTION

In the collections of the American Museum Congo Expedition there
is a single specimen of the African manatee which was secured by Mr.
Herbert Lang at Banana in August, 1915, just previous to his departure
from Africa after six years of collecting in the upper Congo basin. The
specimen, which is now preserved as a tanned skin, cleaned skeleton, and
palatal pads in alcohol (A. M. N. H. No. 53939), is that of an immature
individual. Photographs were made of this animal in the flesh, and three
of them are here reproduced (Plate XX VII). The date of collection is
not definitely established, and there is no record of measurements or of
sex, though the form of the pelvic bones indicates that the animal is a
female.

The occurrence of the manatee in the lower Congo has long been
known, but no skeleton of a specimen from so far south has ever been
described or compared with those from the northern part of the range,
and for this reason the Congo Expedition manatee is of more than usual
interest. As manatees are highly variable mammals and specimens few,
any documented specimen is important, but this individual is particu-
larly so since it is the sole representative of its species in the otherwise
excellent American Museum collection of manatees. Because of the
progressive extermination of manatees and the total inadequacy of
material now in museums of the world, it is unlikely that we shall ever
have a very complete picture of the geographical variation of these
animals. As nearly as I have been able to ascertain, there are no speci-

1Scientific Results of the Congo Expedition. Mammalogy, No. 14.
533

534 Bulletin American Museum of Natural History [Vol. LXVI

mens of manatees either from the upper Niger River or from Lake Chad,
though their occurrence in the former is virtually certain, and their
former occurrence in the latter at least probable.

The most recent reviser of the genus Trichechus, Hartlaub (1886),
assembled more African manatees than any other worker before or since,
but it seems that these were compared with the American coastal mana-
tees from Surinam only, and some of the characters by which Hartlaub
distinguished senegalensis from the group he called latirostris have proved
invalid in the case of skulls from Florida, Puerto Rico, and Guatemala.
Furthermore, his studies were limited to skulls and did not bring out
characters found elsewhere.

This report establishes a few hitherto unrecognized characters for
the distinction of the crania of the different species of manatees, corrects
certain erroneous conclusions made by Hartlaub, and records for the
first time specific features of the postcranial skeleton. A review of the
nomenclatorial history of the manatees is given, and an earlier name than
that currently in use for the African manatees is established as valid;
the type locality for several nominal species is fixed, or restricted, and the
Florida manatee is recognized as a subspecies of the West Indian form.
Literature concerning the range of the African manatee is reviewed in
order to correct conflicting statements that occur occasionally even in
comparatively recent papers.

NOMENCLATURE OF THE MANATEES
TRICHECHUS Linnaeus

1758. Trichechus LiInNaEusS, ‘Systema Naturae,’ 10th Ed., I, p. 34. Type, by
monotypy: Trichechus manatus LinNAEUS. Type locality, by subsequent
designation: ‘“‘West Indies” (Thomas, 1911, Proc. Zool. Soc. London,
p. 1382). By Opinion 112, ‘International Commission on Zoological
Nomenclature’ (Smithsonian Misc. Coll., LX XIII, No. 6, p. 19), Trich-
echus LINNAEUS, 1758, type 7. manatus is placed on the official list of
generic names.

1772. Manatus Brttnnicn, ‘Zoologiae Fundamenta,’ pp. 34, 38. According to
Palmer (1904, p. 398) the type species is Trzchechus manatus LINNAEUS,
1758. Suspension of the Rules in favor of Manatus Briinnich was declined
by the ‘International Commission on Zoological Nomenclature’ (Opinion
A, es Citi) )o

1803. Ozxystomus G. FiscuEer, ‘Das National Museum Naturgesch. zu Paris,’ II,
p. 353. Type: Oxystomus manatus (=Trichechus manatus Linnaeus).

1848. Halipaedisca GistTEL, ‘Naturgesch. Thierreichs f. héhere Schulen,’ p. 83. New
name for Manatus Briinnich, 1772. Type: Manatus americanus.

Date? Monatus D’Orsieny, Keepsake’s ‘Hist. Nat. Desc. Mamm.,’ pp. 256-257,
Pl. xu, fig. 2. Paris. Misprint for Manatus. Original not seen. Refer-
ence from Palmer’s ‘Index Generum Mammalium,’ 1904, p. 398.

1934]

1758.

1784.

1788.

1795.

noo:

1800.

Hatt, African Manatees 539

SPECIFIC AND SUBSPECIFIC NAMES OF RECENT MANATEES
REFERABLE TO T’richechus

Trichechus manatus LINNAEUS, ‘Systema Naturae,’ 10th Ed., I, p. 34.
““HapiraT IN Marr Americano.” The type locality is fixed by Thomas
(1911, Proc. Zool. Soc. London, p. 132) as ‘“‘ West Indies.”

Manati Trichechus BoppAERT, ‘Elenchus Animalium,’ I, p. 173, Rotterdam
(reference from J. A. Allen’s ‘Bibliography of Cetacea and Sirenia.’
Original not seen). According to Allen, this is based on Pennant’s
‘Broad-tailed Manati,’ which was (in the 1781 edition, the only one in
which the name had appeared) applied to both American and African
manatees. For purposes of standardization I assign the type locality
to the West Indies.

Trichechus manatus australis GMELIN, ‘Systema Naturae,’ 13th Ed., I, part
1, p. 60, Lipsiae. ‘Mariafricano et americano.”’ The name was applied
to the true manatees in contradistinction to 7. m. borealis (=Rhytina
borealis). In 1800 its use was restricted by Shaw to the Senegal manatee,
for which another name was already available.

Trichechus antillarum Link, ‘ Beytrige zur Naturgeschichte,’ Band 1, St. 2, p.
109. The name is referred to Buffon’s ‘Grand Lamantin des Antilles,’
for which 7’. manatus was already available. As type locality, I designate
the West Indies.

Trichechus americanus LINK, idem. The name refers to Buffon’s ‘Le Petit
Lamantin de l’Amérique,’ which is a confused account of a small species
of manatee inhabiting American waters from the Amazon to Campéche
and Cuba, including records now referable to the Central American
manatee and the true South American river manatee. I assign the
West Indies as the type locality.

Trichechus senegalensis LINK, idem. Based on Buffon’s ‘Le Petit Lamantin
du Sénégal,’ which in turn was founded on Adanson’s description and the
skull which he secured in Senegal. This latter is said to be in the Paris
Museum. The type locality may be fixed as Senegal.

Manatus aequatorialis LAckPEDE, ‘Tableaux des Mammiféres.’ An. VII
[1799], p. 17, Paris. Manatus aequatorialis is the only species of the
genus named, and since the name is unaccompanied by diagnosis, and
since it is not referable to any one species, it is a nomen nudum.

Manatus Guyannensis BECHSTEIN, 1n Pennant’s ‘Algemeine Uebersicht der
vierfiissigen Thiere,’ II, p. 732. Weimar. This name, was applied
to Pennant’s ‘“‘Guiana manatee,” which is the Trichechus manatus of
Linnaeus.

Manatus Oronocensis BECHSTEIN, idem. This name was applied directly to
Pennant’s “‘Orinoko manatee,’ for which Shaw this same year proposed
the name Amazonius. Despite the name given, I assign the West Indies
as type locality on the ground that Pennant’s “‘species’’ was founded on
Buffon’s composite ‘Le Petit Lamantin de ]’Amérique,’ and any other
assignation would necessitate revision of nomenclature or lead to con-
fusion.

536

1800.

1802.1

1815.

1816.

1824.

1838.

1848.

Bulletin American Museum of Natural History [Vol. LXVI

Manatus stroggylonurus BECHSTEIN, idem. The name was proposed for Pen-
nant’s ‘“‘Round-tailed manatee,” for which Link’s name is already avail-
able.

Trichechus Clusiw SuHaw, ‘General Zoology,’ I, part 1, p. 246. Based on Clu-
sius’s figure and description of a West Indian manatee, therefore a
synonym of T'richechus manatus Linnaeus. Its type locality may be
given as the West Indies.

Trichechus A mazonius SHAw, ‘General Zoology,’ loc. cit. Based on the reports
of manatees in South American rivers, and referred to Pennant’s ‘‘Orino-
ko manatee.’’ Pennant specified that this was Buffon’s ‘Le Petit
Lamantin de |’Amérique,’ but Buffon, Pennant, and Shaw failed to
define their animal clearly and confused the West Indian and the true
river manatees. By reason of the foundation of the species, and despite
the name, I designate the type locality as ‘‘ West Indies,” so that current
nomenclature may not be unnecessarily disturbed.

Manatus minor Davptn, ‘Histoire Naturelle’ of Buffon, Didot Edition,
‘Quadrupeds,’ XIV, p. 194. Stated to be ‘Le Petit Lamantin d’Amér-
ique’ and correctly referred to the account in Vol. IX, p. 251.
This name has then the same status as americanus Link. I hereby fix the
type locality as ‘‘ West Indies.”

Manatus fluviatilis ILLigER, Abhandl. d. Kon. Akad. d. Wissens. in Berlin,
1809-1811, p. 110. This name, given without diagnosis, appears with M.
americanus in a list of South American mammals and is a nomen nudum.

Manatus sphaerurus ILLIGER, op. cit., p. 79. The name appears together with
Halicore cetacea in a list of African mammals. No diagnosis is given it,
and it is therefore a nomen nudum.

T |richechus|, M|anatus| africanus OxEn, ‘Lehrbuch der Naturgesch.,’ Th. III,
Abt. II, p. 688. A short diagnosis is given, and the presence of the
species in Senegal and the Congo is noted. The type locality I here
designate as Senegal.

Manatus latirostris HARLAN, Journ. Acad. Sci., Philadelphia, III, p. 394.
The name was tentatively proposed for the manatee from the coast of
eastern Florida if this should later prove distinct from senegalensis. A
description based on two skulls is given. Presumably these are in the
museum of the Philadelphia Academy of Natural Sciences.

Manatus atlanticus OxEn, ‘Allgemeine Naturgeschichte,’ Abt. II, Band VII,
p. 1098. The name is applied to both American and African manatees,
followed immediately by the name T'richechus manatus. I propose the
West Indies as its type locality.

Manatus nasutus WYMAN, Proc. Boston Soc. Nat. Hist., II, p. 199. The name
is proposed in a footnote (signed ‘‘J. W.’’) to an article by G. A. Perkins,
describing a specimen from the Caracalla River, twenty miles east of
Cape Palmas, Ivory Coast. The specimen on which the original descrip-
tion is assumed to have been based is at present in the Museum of
Comparative Zodlogy.

1For the establishment of 1802 as the publication date of Lacépéde’s and Daudin’s *Tableaux,’
see C. Davies Sherborn, 1899, ‘Natural Science,’ XV, pp. 406-409.

1934] Hatt, African Manatees 537

1856. Manatus Vogeliti OwEN, Edinburgh Phil. Journ., N.S., IV, p. 346. The name
was tentatively proposed for the specimen taken by Vogel’ from the
river Benue.

1861. Manatus Owenit Du Cuattuu, Proc. Boston Soc. Nat. Hist., VII, p. 367.
Type locality: the Camma country, in the mouth of the Gaboon. The
British Museum has four of the Du Chaillu specimens, the Mus. Coll.
Surgeons, one.

1883. Manatus inunguis NATTERER (in Pelzeln), Verh. Zool. Bot. Ges. Beiheft,
XXXII, p. 89. Pelzeln, who quoted Natterer’s notes in full, considered
inunguis a synonym of australis. Natterer’s name is based on five speci-
mens from the Rio Madeira, Brazil. Though several other older names
have been used for this manatee, none of these names is clearly restricted
to this one species.

1897. Manatus Koellikeri KUKENTHAL, Zool. Anz., XX, p. 40. Type locality:
Surinam and hence a synonym of manatus. No specimen is mentioned.

The African Manatee

Trichechus senegalensis Link

Phoca manatus Brisson, 1762 (part), ‘Regnum Animale,’ p. 164. This is a re-
combination of Linnaeus’s T’richechus manatus, but includes the African manatee.

Manat: Trichechus BopDAERT, 1784 (part) ‘Elenchus Animalium,’ I, p. 173,
Rotterdam. ‘Reference from Allen’s ‘Bibliography.’ Original not seen.

Trichechus manatus australis GMELIN, 1788 (part), ‘Systema Naturae,’ 13th Ed.,
I, part 1, p. 60, Lipsiae.

Manatus australis Rerzius, 1794 (part), Kungl. Sven. Vet. Akad. Handl., XV,
p. 291.

Trichechus senegalensis LINK, 1795, ‘Beytrige zur Naturgeschichte,’ Band I,
St. 2, p. 109. The name is referred to Buffon wherein is described (1782, Suppl., VI,
p. 403) ‘Le Petit Lamantin du Sénégal,’ based mainly on Adanson’s account. Adan-
son’s Senegal skull in the Museum d’ Histoire Naturelle, Paris, is described in Buffon
(1765, XIII, pp. 431-432).

Trichechus aequatorialis LACKPEDE, 1799 (part), ‘Tableaux des Mammiféres,’
An. VII [1799], p. 17, Paris.

Trichechus Australis SHAw, 1800, ‘General Zoology,’ I, part 1, p. 244. Thisname
is here limited to the African manatee. A meager description is given, and a specimen
inthe Leverian Museum noted. Pennant’s fanciful figure of a “‘round-tailed
manatee”’ is reproduced.

Manatus stroggylonurus BECHSTEIN, 1800, in Pennant’s ‘Algemeine Uebersicht
der vierfiissigen Thiere,’ II, p. 732. Weimar.

Trichechus senegalensis DAaupDIN, 1802, in Lacépéde’s and Daudin’s ‘Tableaux des
Mammiféres’ in Buffon’s ‘Histoire Naturelle,’ XIX, ‘Quadrupeds,’ XIV, Didot and
Didot Edition, p. 194. Said to be ‘Le Petit Lamantin du Sénégal’ and referred to
Vol. IX, p. 254, where a description of the Senegalese manatee taken chiefly from
Adanson is found.

1This communication was read before the 1856 meeting of the British Association for the Advance-
ment of Science. It was first published in the Edinburgh Philosophical Journal and subsequently
(1857) with slight differences in the Report of the 26th Meeting of the British Assoc. Adv. Sci. and in
the “Institute” (1857). I have not seen the latter reference.

038 Bulletin American Museum of Natural History [Vol. LXVI

Manatus sphaerurus IuLticmR, 1815, Abhandl. d. Kon. Akad. d. Wissens. in
Berlin, 1809-1811, p. 79.

Trichechus, Manatus, africanus OkEN, 1816, ‘Lehrbuch der Naturgesch.,’ Th.
III, Abt. II, p. 688.

Manatus senegalensis DESMAREST, 1817, ‘Nouv. Dict. Hist. Nat.,’ XVII, p. 262.
Based on Cuvier’s (1809, Ann. Museum, XIII, pp. 294-296, Pl. x1rx, figs. 4-5)
description and figures. ‘Though currently accepted as the correct name of the
African manatee, Desmarest’s name must be considered a homonym of Link’s.

Manatus atlanticus OKEN, 1838 (part), ‘Allgemeine Naturgeschichte,’ Abt. II,
Band VII, p. 1098.

Manatus nasutus WYMAN, 1848, Proc. Boston Soc. Nat. Hist., II, p. 199.

Manatus Vogelat1 OWEN, 1856, Edinburgh Phil. Journ., N.S., IV, p. 346.

Manatus Owent Du CHaiLuu, 1861, Proc. Boston Soc. Nat. Hist., VII, p. 367.

It is convenient to state at the outset my conclusion that Hartlaub
was quite right in the recognition of but three full species of manatees,
though geographical representatives may, in some cases, be worthy of

subspecific recognition. The correct names for these manatees are:

Trichechus manatus manatus Linnaeus

RancEe.—The West Indies, the borders of the Caribbean Sea, the
coast and lower reaches of the rivers of northeastern South America.

Trichechus manatus latirostris (Harlan)
Rance.—The coast and coastal rivers of the United States from
Beaufort, North Carolina, to the Florida Keys and the coasts of the
Gulf of Mexico.

Trichechus senegalensis Link
RanGu.—The west coast and coastal rivers of West Africa from the
Senegal to the Quanza, the upper reaches of the Niger, and probably
the Lake Chad drainage. (See pages 554 to 560.)

Trichechus inunguis (Natterer)

Rance.—The rivers of northeastern South America, particularly
the Amazon and Orinoco systems.

The only changes from the nomenclature applied by most modern
writers are the replacement of T’richechus senegalensis Desmarest, 1817,
by Trichechus senegalensis Link, 1795, and the adoption of latirostris
as a valid race of manatus instead of as a synonym of manatus or the
name of a species distinct from the latter.

1934] Hatt, African Manatees 039

SPECIMENS EXAMINED

In the course of this study I examined the collection of manatees in
the American Museum, which, at the time, consisted of the following:
T. senegalensis
1 skin, skeleton, Congo River
T. m. manatus
1 skeleton, Puerto Rico
1 skull, Honduras
T. m. latirostris
9 skeletons, Florida
2 skulls, locality unknown
T. inunguis
2 skeletons, Amazon
15 skulls, Amazon
1 embalmed specimen, Amazon
1 cast, locality unknown.
I also studied the following material, generously loaned by the Field
Museum, to the officers of which I am greatly indebted.
T.m. manatus
4 skulls, Guatemala
T. m. latirostris
1 skull and partial skeleton, Florida
1 skull and partial skeleton, Texas (?)
T. inunguis
2 skulls, ‘“‘Para.’’ (Probably from farther up the Amazon.)
Furthermore I had the carefully executed figures of Hartlaub,

Blainville, and certain others for comparison.

CRITERIA FOR THE CLASSIFICATION OF THE MANATEES

Material available for diagnosis of the external features of manatees
is limited because of the inadequacy of published descriptions and the
paucity of embalmed specimens, casts, reliable drawings, and good
photographs. ‘Tanned skins are uncommon and, when accessible, are so
shapeless and changed as to be all but useless. Observations on the
internal soft anatomy, though of great generic interest, are not of
demonstrated value for specific diagnosis.

The skeletons have yielded much the most important and reliable
information as to the differences in these animals, and as always, the
skulls have been more commonly preserved and studied than other bony
parts. The postcranial skeleton does, however, show variation that is
correlated with age, sex, and specific habitus.

540 Bulletin American Museum of Natural History [Vol. LXVI

INDIVIDUAL VARIATION AND ASYMMETRY

The range of variation in manatees is so extensive that Gray de-
spaired of finding stable characters on which species nomenclature could
be founded. Better series of specimens than were available to Gray,
however, allow some sorting out of the characters, and it is an easy
matter to distinguish at least three species of manatees, no matter how
extensive their peculiarities, though occasional specimens are encountered
that vary, not only in the form and proportion of a few characters, but
very markedly in almost every character. Lacking a good series for
comparison, these variants would appear to represent well-marked
‘““species.”’ Thus I examined the skeleton of one medium-sized Florida
manatee in which the bones are lighter, more compact, and more strongly
ridged than in any other specimen.

Asymmetry is uncommon and where present is pathological. On
one young Florida skull the left occipital condyle is very nearly double
the size of the right, but the bone forming it is rough and irregular. In
another Florida specimen the diameters of the humerus, radius, and ulna
of the left side are approximately 25 per cent greater than those of the
other side, but here too the form of the bones is definitely anomalous.
The sternum of a third Florida specimen is greatly warped to one side.
The teeth of an adult Amazonian animal are completely disorientated,
some of the teeth of each row, even those unerupted, lying with the
direction of their ridges as much as 180° from the normal position.

THe EXTERNAL ANATOMY

When the wide range of variability shown in the external form of the
species manatus, as documented by several accurate drawings, good
photographs, and casts, is considered, it is deemed that from external
characters senegalensis and manatus cannot be told apart. However,
the species 7nunguis is, in all likelihood, constantly characterized by the
absence of nails, a white breast patch, slender proportions, and elongated
flippers.

THe VERTEBRAL COLUMN

The Congo specimen possesses six cervical vertebrae, as do the
other members of the genus. The rib-bearing vertebrae vary in number
in Florida manatees (in a series of 11 skeletons) from 17 to 19; a Puerto
Rican specimen has 17 such vertebrae, and each of two Amazonian.
manatees has 15 pairs of ribs. (Blainville’s figure shows 16 pairs.)
The presence of 17 pairs in the Congo specimen (Biittikofer’s specimen

1934] Hatt, African Manatees 041

bore 18 pairs of ribs according to Jentink, 1888, p. 33) seems then to be
in line with this form, being more nearly allied to the species manatus
than inunguis. The numbers of lumbo-caudal vertebrae vary in the
same Florida specimens from 27 to 29 (without respect to the number
of rib-bearing elements in the same skeleton). In the Puerto Rican and
Congo specimens there are 25 lumbo-caudals. Twenty-six were found in
Bittikofer’s Liberian manatee (Jentink, loc. cit). In the two Amazonian
specimens there are 25 (older) and 22 (younger).

Other than this variation in numbers, in which specific tendencies
seem to be weakly manifest, no characters of the column show any
constant differences between the forms.

THE STERNUM

The sterna of manatees present well-marked shapes that, though
individually variable, are specifically constant in certain features.

The sternum of senegalensis is much like that of manatus, except
that the African species does not (in the Congo specimen ) have a deep
median notch in the anterior border of the bone but may have two
light notches flanking a median prominence. This difference may be
clearly seen. by an examination of figure 1. The margin of the caudal
prolongation of the sternum in the Congo specimen is also incised, a
condition uncommon in the species manatus.

The sternum of znunguis is a smaller bone in proportion to the
size of the animal than that of manatus and senegalensis and may be
recognized by its slender proportions and backwardly directed lateral
processes.

THe APPENDICULAR SKELETON

THE PEcTORAL GIRDLE.—The pectoral girdles of manatees differ in
the proportions of their component parts, but there is no deviation from
the essential form. As in most other characters, the South American river
manatees differ most widely from the type species manatus, for their
flippers are proportionately longer than those of the other members of
the genus, a lengthening which has occurred chiefly in the metacarpals
and phalanges.

The scapula (Fig. 2) of the Congo manatee more closely resembles
that of znunguis than that of manatus. From both it differs in being
comparatively long and narrow and in having the coracoid border gently
curved and without a pronounced angle at the coraco-vertebral juncture
or above the incisura. The spine is apparently a little higher, the

Fig. 1. Sterna of manatees.

a, T. manatus latirostris, Florida, A.M.N.H. No. 91096; b, T. m. latirostris, Florida, A.M.N.H.
No. 35219; c, T. senegalensis, Congo, A.M.N.H. No. 53939; d, T. inunguis, Amazon, A.M.N.H. No.
94163; e, T. inunguis, Amazon, A.M.N.H. No. 94164.

Fig. 2. Scapulae of manatees.

a, T. inunguis, Amazon, A.M.N.H. No. 94163; 6b, 7. senegalensis, Congo, A.M.N.H. No. 53939;
S T. manatus latirostris, Florida, A.M.N.H. No. 90178; d, T. m. manatus, Puerto Rico, A.M.N.H. No.
35566.

544 Bulletin American Museum of Natural History [Vol. LXVI

acromion thinner, the tuberosity of the spine less pronounced. In the
matter of proportionate narrowness, znunguis is intermediate between
senegalensis and manatus, but in other characters it can hardly be
distinguished from manatus. The geographical representatives of
manatus show no constant peculiarities of the scapulae.

The humerus of the Congo Expedition manatee is distinctly thinner
than that of Florida manatees of the same length. The weight of the
Congo humeri is just half that of a typical pair of the same length from
Florida. In this respect the Congo manatee is very closely similar to
inunguts.

The relative proportions of the humerus are carried into the radius
and ulna. In both senegalensis and inunguis these two elements are
about one half the diameter of corresponding elements in manatus.

The metacarpals of the three species differ markedly in length.
The fourth digit is the longest, and measurements were therefore made
on the metacarpal and phalanges of this finger of each specimen. This
metacarpal grows more rapidly or for a longer period than does the
radius, and in consequence the metacarpal is proportionately longer in
older specimens than in young, when, as here, the length of the radius is
used as the unit of comparison. In specimens of approximately equal
age, however, differences are marked. Thus in manatees of the approxi-
mate size of our Congo specimen the metacarpal is about 54 per cent of
the radial length in manatus, about 64 per cent of the radial length in
inunguis, while in the Congo specimen the metacarpal is 62 per cent as
long as the radius.

In the matter of the length of the first carpal, senegalensis is not
intermediate between the Caribbean manatees and znunguis but falls
within the limits of variation of manatus. This first phalanx of the fourth
digit does not grow as rapidly or over as long a period as its metacarpal,
with the result that with increased age it is in proportion to this meta-
carpal increasingly shorter, whereas its growth rate outstrips that of the
humerus, and increased age brings a proportionately, as well as an
actually increased length.

Again comparing immature animals about the size of the Congo
specimen, it is found that the first phalanx in both manatus and sene-
galensis is about 25 per cent of the humerus length, or 40 per cent of the
metacarpal length (the specimens suggested that this element in the
Florida manatees is considerably longer than that in the Puerto Rican),
whereas in :nunguzs the corresponding percentages are 40 and 60.

The remaining phalanges follow the trend indicated by the first,

1934] Hatt, African Manatees 545

senegalensis and manatus being essentially short-fingered and znunguis
long-fingered.

Tue Prtvic GirpLE.—The immaturity of our Congo manatee
precludes the determination of specific characters which might appear in
the innominate bones, but these bones are sufficiently developed to
furnish a key to the sex of the specimen and show it to be a female.
These bones are indistinguishable from those of immature female Florida
manatees and from those of certain Surinam specimens figured by Krauss.
My failure to find any innominate bones in two uncleaned inunguis
skeletons fresh from the field, and the lack of mention or figuration of
these bones in literature may indicate that this species has completely
lost its pelvic girdle.

THE SKULL

A glance serves to identify the skull of Trichechus inunguis, but the
differences between the skulls of senegalenszs and manatus are more
subtle. The two manatees last mentioned differ from znunguzs, not only
in the general shape of the skulls, but in the very character of the bone
itself, for though the skull bones of senegalensis and manatus are in general
dense and smooth, those of ¢nwngwis are, with rare exceptions, soft,
chalky, and rather elaborately roughened. The skull bones of this latter
species are also lighter in proportion than those of the other two.

The general skull shape of senegalensis and manatus is broad and
compact and the snout short (senegalensis being shorter than manatus
in this respect), whereas the skull of znunguzs is lengthened and char-
acterized by a long snout. The conspicuous recession of the nasals and
the posterior border of the anterior nasal opening leaves at the anterior
end of the skull the characteristic large nasal basin, the floor of which is
formed by the palatal parts of the maxillae and the vomer. The form of
this basin is dependent in the main upon the extentof the forward growth
of the premaxillae, and this is a process progressing with age to maturity,
or possibly throughout life; but when comparisons are made of skulls of
equal length, specific patterns are discernible. This nasal basin is
broader in the adult African manatee than the corresponding area in
manatus, but the difference is very slight, and in immature specimens it is
impossible to distinguish these two species on the basis of this feature.
The adult manatees of the western Caribbean and Gulf of Mexico appear
to be slightly longer snouted than corresponding specimens from Florida,
but additional material would be necessary to establish the constancy of
this probable difference. The nasal basin of the Amazonian manatee is,

546 Bulletin American Museum of Natural History [Vol. LXVI

however, so very distinct that the smallest skulls show the character-
istic long, narrow basin of this species.

The nasal process of the premaxilla of senegalensis covers the
anterior part of the nasal cavity surface of the orbital process of the
frontal to some extent, whereas in znwnguis and manatus there is little
or no such covering, the upper border of the premaxilla lying below the
lower border of the frontal. This suture in senegalensis also differs from
the other species in being distinctly shorter, a configuration which is
accompanied by a relative rounding off of this end of the bone as con-
trasted with the sharply pointed ends found in adult manatus and inun-
guis. The specimen from the Congo does not, however, fit in with
Hartlaub’s characterization of senegalensis as a manatee with the nasal
processes of the premaxillae distally expanded, for in this specimen these
processes show less terminal expansion than some Florida specimens of
the same size.

The anterior frontal margin between the roots of the orbital process
in the African manatee is typically smooth and unserrated, but in this
character it does not diverge strongly from manatus in which, particularly
in old individuals, the frontal margin is a little jagged. The breadth of
this margin is, however, considerably greater in manatus than in sene-
galensis, and the combination of relative breadth and irregularity or
smoothness of this margin would probably suffice to identify any
specimen.

The nasalia of manatees have been made known by Krauss and
Hartlaub, but the statements concerning them rest on an insecure footing
through the extreme variability in these bones and their frequent loss
in the course of museum preparation. Hartlaub concluded that the
senegalensis nasalia were typically platelike, that gradations to peglike
bones were not infrequent, and that it was possible that in some animals
these bones were completely undeveloped. The Congo Expedition
manatee possesses a very small and spongy left nasal bone, lightly fixed
in a small pit of the frontal. The nasal does not come nearer than 8
mm. to the ascending branch of the maxillary. There is but slight indica-
tion that the right nasal bone was ever present, though of course it may
have been present and free.

The nasalia of manatus, as Hartlaub found, are typically thick
almond-shaped bodies, but his small series of four specimens does not
represent the range of variation, for I have found in a better series varia-
tion from large peglike bones to platelike types and have noted that in
some cases the nasalia do not seem to have been developed. Twelve

1934] Hatt, African Manatees 547

skulls of Florida manatees examined range from 255 to 375 mm. in total
length, or from youth to full maturity. In these specimens there is a
great range of nasalia, from apparent total absence to well-developed peg-
shaped bones lying in deep sockets of the frontal and uniting in a loose
suture with the ascending rami of the maxillae, and (or) the premaxillae.
In length these vary from 20 to 48 mm. in adult specimens. Of two skulls
identical in size, one has large, long nasal bones, the other bears no
evidence that nasal bones have ever been present.

A large manatee, reputedly from Texas, has nasalia thoroughly
typical of those from Florida. Four specimens from Guatemala present
extreme variation, two having peg-shaped bones, one with flat nasalia
horizontally placed and roofing the posterior part of the nasal trough, and
the fourth with its pair of nasalia flat and vertically placed as are
those in specimens of znunguis. A Puerto Rican manatee differs in no
respect from certain Florida specimens in regard to its nasalia.

The skulls of zrnunguis examined ranged in greatest length from 200
to 360 mm., from early youth to maturity. Ten of these were cleaned,
three had been roughed out in such a way that the nasal bones, if present,
may have been lost in the field, and six were roughed out so that there
was very little possibility of the nasal bones having been lost. I cut away
the dried flesh of these six skulls in a search for nasal bones with the
following results: In two skulls, 265 and 330 mm., in greatest length,
there was no trace of nasalia, either actually, in pits for their reception, or
in grooved surfaces for articulation. Neither was there any indication
that nasals had fused to the frontal bones. In another specimen, 355
mm. long, there was no trace of a nasal unless a minute conical nodule (4
mm. long) lying in the site of a right nasal was a vestige of this bone. On
the right side of a fourth specimen, 270 mm. long, there was a well-
developed plate of very compact bone lying totally free, but closely asso-
ciated with the vertical surface of the frontal bone at the anterosuperior
border of the nasal chamber. There was no trace of a corresponding bone
on the opposite side. The two remaining skulls, 260 and 330 mm. long,
bore nasalia on each side similar to that mentioned. The five such nasalia
examined were alike in being of a modified lozenge-shape (triangular or
quadrangular), though their greatest diameters ranged from 25 to 13
mm., their minima from 15 to 11 mm., and their thicknesses from 3 to 6
mm. These bones lay with their long axes anteroposterior, and their
chief plane vertical. They were in loose contact with the median free
surface of the orbital branch of the frontal directly posterior and ventral
to the anterior median edge of the frontal. In one case the lateral

548 Bulletin American Museum of Natural History [Vol. LXVI

branch of the ethmoturbinals bounded the bones medially. The surface
of the frontals with which the nasals were in proximity was in each case
slightly concave in accommodation to these bones.

The lacrimal bone of the right side of the Congo skull is in place,
but that of the opposite side is missing, due, obviously, to over-macera-
tion. Since Hartlaub found these bones preserved in only one of the ten
African skulls examined, and this in a newborn specimen, it appears
worth while to note the condition of the lacrimal remaining in the Congo
skull. It is, in general, intermediate in size, position, and shape be-
tween the type characteristic of nungwis and that invariably occurring
in manatus. The bone is at its broadest exposed point 3 mm. in thickness
and appears to taper down into its groove between two laminae of the
maxillae. Its upper border is nearly in contact with the orbital process of
the frontal, while its lower edge touches the edge of the jugal. A tri-
angular surface is freely exposed laterally and forms part of the antero-
medial wall of the orbital ring.

The lacrimalia of the znwnguis, which I examined closely, resemble
those described by Hartlaub. In each case their scalelike nature is
evident, and they are quite unlike those of any manatus which I have
examined. In most of the manatus studied these bones are in place, and
in each case they are essentially similar, large, thick, and more like the
same bones in senegalensis than in znunguis. Though the lacrimals are
often missing, it is very clear that they had been present, and large, in
every manatus skull which I saw.

The vomers of manatees show strong specific characters in their
length. In senegalensis they are always short, extending approximately to
the level of the middle of the orbit. In manatus they are long and,
except in the newborn, reach to the foramen incisivum or beyond. In
inunguis the vomer is intermediate in length between that of senegalensis
and manatus, being in the newborn very nearly as short as in the African
species, and in old specimens occasionally reaching to within an inch of
the incisive foramen.

Low edges appear on either side of the floor of the nasal chamber
anterior to the orbital region. In senegalensis these edges are probably
constant. Though Hartlaub concluded from an examination of Surinam
specimens that there were no such ledges in manatus, I find them well
developed in most, but not all, of the specimens at hand. These ledges
are usually present in inunguws but are lightly developed and farther to
the rear than in the other species.

The circumorbital region of the manatees shows something of a

1934] Hatt, African Manatees 549

different pattern in the three species. In senegalensis the orbital process
of the frontal diverges most strongly in a lateral direction, and the upper
orbital borders formed by this process are strongly convergent in the
African and Amazonian manatees. This latter character is a configura-
tion well marked even in the newborn. If the line of the outer border of -
this orbital process is extended forward, it will cross the median line
anterior to the end of the skull in manatus, whereas in senegalensis and
inunguis this crossing occurs within the limits of the skull.

The age of the individual manatee determines the degree of back-
ward divergence of the postorbital process of the orbital process of the
frontal, and at the same time the closure of the orbital ring. In all
species this occurs only late in life and is seen in few specimens, though
it would appear that this closure is more frequent and occurs earlier in
life among Guatemalan than among Florida manatees.

The infraorbital foramen in manatees is ordinarily simple, though it
may be divided in manatus. Hartlaub observed that in this species
the division was frequent, but, with the exception of two of four Guate-
malan skulls examined, I have not found such division in any specimen
of manatus. As most of my manatus skulls are from the northern limits
of the species, and those of Hartlaub from the southern limits, it is
possible that a divided infraorbital foramen is of more frequent occur-
rence in the south than in the north.

The bony ridges formed at the superior border of the temporal
muscle are more or less vertically directed in senegalensis and manatus,
while in znunguis these ridges are produced laterally and do not rise
above the general level of the skull roof.

The malar process of the temporal in manatees is swollen and spongy
in nature. At the anterior end the thin layer of compact bone sheathing
the spongy mass is frequently discontinuous, and a rough or perforated
surface is common. However, when the surface is reasonably uninter-
rupted, it is smooth in senegalensis and always rugose in inunguis. This
same area in manatus from Florida is fairly well-grooved in most speci-
mens, though this condition does not appear to apply to the species as a
whole, the same area in a manatee from Puerto Rico, one from Texas (?),
and three from Guatemala being smooth.

The zygomatic process at its base is much thicker in senegalensis and
manatus than in inunguis.

The malar bone sends a process downward which, in senegalensis, is
broad and sharply truncated. Specimens of manatus closely approach
this shape in some instances, but often have a backwardly directed

900 Bulletin American Museum of Natural History [Vol. LXVI

process, in this respect resembling inunguis, although the ventral malar
process in 7znunguts is always narrow and sharply tipped.

The supraoccipital bones of senegalensis and manatus are of a com-
mon pattern but differ very strongly from the same bones in the Ama-
zonian manatee. ‘The former present a very nearly flat, transverse plane,
whereas the latter are rugose and so mounded that the lambdoidal ridge
as seen from above is yoke-shaped.

The outer borders of the exoccipitals of senegalensis are knotty,
pitted, and rough. In this respect the African species is similar to most
munguis and unlike manatus.

The foramen magnum of senegalensis and inunguis is roundish,
while in manatus it is oval. This shape is modified in part by differences
in the dorsal rim and also, as seen from the ventral surface, by the notch-
ing of the lower border. In manatees from the Congo, the Amazon,
Puerto Rico, and Guatemala, that is to say, in representatives of all
species, the dorsal border is strongly curved. Florida and Texas speci-
mens of manatus, however, have flat dorsal rims, perhaps one of the most
constant features on which the nominal race latirostris may lay claim to
subspecific individuality from manatus. ‘The notching of the lower
border (the basioccipital) is also a fairly constant and peculiar mark of
the northern representatives of the species manatus. These differences in
the limits of the foramen may be partially synthesized by taking the
ratio of greatest vertical diameter to greatest horizontal diameter. Ex-
tremes are found in a specimen from the Amazon in which the vertical
diameter is 75 per cent of the horizontal, and in a Florida specimen in
which the corresponding percentage is but 54 per cent. There are no
specific limits to these ratios, however, for the Congo skull has a foram-
inate index of 0.66; skulls of znwnguis vary from about 0.65 to 0.75; and
specimens of manatus from 0.54 to 0.71. Within the species, however,
there seem to be geographic trends, for Florida specimens have indices
of from 0.54 to 0.61; a Texas specimen an index of 0.60; while four
Guatemalan manatees have corresponding indices ranging from 0.66
to 0.71; anda Puerto Rican manatee has an index of something over 0.70.

Basally, skulls of the species of manatee may be recognized by a
number of characters. Over the occipitosphenoidal suture is an
eminence which in inunguis is median and simple, while in the other
species it is lateral and double, and anterior to this the shape of the
posterior nares is distinctive, though variable with age. In the Amazoni-
an manatees the opening is sagittate in young individuals, while in old
manatees it broadens out ventrally so that it is bicordate or presents the

1934| Hatt, African Manatees dol

form of a double, symmetrically notched circle. The posterior nares of
senegalensis are very nearly circular, whereas those of manatus are deltoid.

The pterygoid process of the manatees is formed by wings of the
alasphenoid, palatine, and pterygoid bones, and this compound process
usually ends distally in three more or less distinct points that are
aligned in a sequence of lateral, intermediate, and medial, best seen from
the rear. These points correspond fairly closely to the distal ends of the
three bones listed and may be called alasphenoid, palatal, and pterygoid
points, respectively. In the species inunguzs the pterygoid process is
long and narrow, and the palatal point is much the highest of the three.
In the Congo manatee the pterygoid process is also long and narrow, but
here the palatine and pterygoid points are coequal and longer than the
alasphenoid. No general rule applies for all specimens of the species
manatus that I have examined. In the Florida and Texas specimens
that I have seen the pterygoid process is similar to that of the Congo
manatee, except that it is usually thicker. In manatees from Guatemala
and Puerto Rico, however, the pterygoid process is short and very broad,
with either the pterygoid, or palatine point longer.

According to Hartlaub, the foramen incisivum is always simple in
senegalensis, a condition also found in the Congo specimen. The same
author observes that this foramen is often completely or incompletely
divided in znunguwis. The foramen of manatus, however, may not be
described as simple, for in a large percentage of the specimens that I
have examined there is a partial division into anterior and posterior
incisive foramina. Although the division is not complete in any case
seen, it is sometimes nearly so.

The anterior end of the palate of the manatees is covered with a
heavy plate that leaves a roughened area on the under side of the pre-
maxillae and maxillae. This surface is broadest at about the juncture
of the two bones and is constricted just anterior to the level of
the foremost teeth. This constriction is most pronounced in inunguis,
the width at the constriction being usually about one half that at the
maxillary-premaxillary suture. The constriction in manatus and
senegalensis, though individually variable, is usually not great, the least
width being about 85 per cent of the greatest.

The molars! of senegalensis and manatus seem to me to be indistin-
guishable, though they differ strongly from those of inunguis. These

1At the time the manuscript of this report was in the hands of the printers, I saw in the British
Museum a skull of 7. manatus (B. M.370E) from Surinam with a left upper incisor in place. Thisskull
had a greatest length of 325 mm., so was adult. The incisor, fattened and almost straight-sided,
measured 15.5 mm. in length. Its tip was well worn. There was a single root, but the tooth was per-
forated by a foramen from the center of its anterior to the center of its posterior face.

Divs Bulletin American Museum of Natural History [Vol. LXVI

latter are of a smaller diameter and are strongly furrowed. In all
of the species there are anterior and posterior cingula, but though these are
smooth in the unerupted teeth of manatus and senegalensis, they are
deeply furrowed in znunguzs. In the first two species each of the two
great transverse ridges is divided into three cones or sectors, but in the
Amazonian manatee these primary cones are more or less broken up
into a series of other smaller cones. |

THE MANDIBLE

Hartlaub’s analysis of the characters of the mandible of senegalensis
is good, except that some of the features he assigned to the mandible
of znunguzs and manatus do not well apply to the series of these species
that I have examined, and a few minor differences appear to have
escaped his notice.

The interramal interval is broad in senegalensis as it is in manatus,
a feature by which these species are again easily distinguished from 7n-
ungurs, and the rami of the latter species lie more nearly in parallel planes
than is the case in the other two species. The African species, however,
differs from manatus in that there is less of a constriction in the diameter
of the ventral border between the body and the angular process. In
senegalensis this process is wider and more in line with the ramus than
in the other species.

The symphysial suture closes early in the African manatee, as
Hartlaub pointed out, and in this species there is no deep furrow along
the anterior margin. In this respect it differs from American manatees.
This furrow, which is most conspicuous in manatus, is a character best
developed at maturity and is not well marked in the newborn or very
young. .

The interior mental fossa is always deeper in young manatees than
in old, but comparing equal-aged material the fossae of African and
Amazonian manatees are, in most instances, deeper than those of Carib-
bean specimens. The character is not constant, however, and, contrary
to the conclusion of Hartlaub, is of little use to the taxonomist.

The anterior end of the mandible of all species is essentially similar
in respect to form, as seen in norma dorsalis. Hartlaub claims that this
region is truncated in senegalensis and inunguis, whereas in the Surinam
manatee (manatus) it is tapered or tipped; but my good series of manatus
shows no difference in this character from the other species, and I would
describe the anterior end of the mandible of manatus as truncated.
However, there is very often a sharp median cone of compact bone ex-

1934] Hatt, African Manatees 553

tending forward from the most anterior part of the symphysis, and though
this spine does occasionally occur in znunguzs, and possibly in senegalensis,
it is almost a constant feature of the mandible of the postnatal manatus.

While not constant either for a species or, probably, even during the
life of an individual, the mental foramina of the manatees follow different
tendencies in inunguis than they do in manatus and possibly show a
slightly different average in senegalensis than in manatus. In wnunguis
these foramina range in number from 11 to 15 (average 15—). The range
in manatus is from 4 to 7 (average 6). In the Congo specimen there are 4
mental foramina, and in Blainville’s figure 3 are shown on one side.

As Hartlaub noted, the ventral border of the mandibular ramus,
presents a greater curve to the horizontal plane in manatus than it does
in the other species; in the others there is no great difference. This
curve results from the deepening of the mandibular symphysis and is
so excessive in manatus that it is one of the most pronounced features of
the species.

The coronoid process of the manatees presents the usual high varia-
tion in shape and cannot be relied upon in diagnosis. Hartlaub concludes
that the posterior superior angle in znwnguis is constantly hooked and
that these hooks are only occasional in manatus. In my series I find that
even in old animals the hooks are not always developed in inunguwis,
though they are usual in the Caribbean manatees. The extent of broad-
ening of the coronoid, which Hartlaub regards as a distinguishing charac-
ter of the species, appears to me to be very weak and unreliable for separa-
tion of senegalensis and manatus material, since in this feature Florida
specimens are indistinguishable from the Congo animal.

The transverse breadth of the condyle of senegalensis is slight com-
pared with that of manatus and inunguis, but the character is so relative
that, lacking a large series of specimens, the difference is not discernible.

The mandibular foramen is subject to certain specific modifications.
Thus, in the Congo specimen it is separated from the bony sheath by a
septum of bone, whereas in znwnguis and manatus no such septa occur,
with the exception that there are, rarely, converging processes that
may nearly touch to form a partial septum. This bony ridge is well
developed in the Congo specimen, and is shown in a figure (Plate 11,
figure 27) accompanying Hartlaub’s paper. However, it does not appear
in the figure reproduced in Flower’s ‘Osteology’ (1870, Fig. 64, p. 197)
and may not be very constant.

A single specimen supposed to have come from Texas is different
from other manatees examined in that the lamina lying medial to the

554 Bulletin American Museum of Naturel History [Vol. XVI

dental trough has expanded so as to mask completely the germinal
wrapper from behind. Whether this is a geographical or an individual
variation cannot be determined without additional material from the
region.

The tooth rows reach farther forward in old age than in youth, but
those of znunguis, unlike the other species, never closely approach the
mandibular symphysis.

THe Hyor ArcH

The hyoid bones of the Congo specimen closely resemble those of
similar aged material from Florida, and it is believed that no characters
for diagnosing the species are to be found in this structure.

THE DISTRIBUTION OF TRICHECHUS IN THE AFRICAN REGION

As far as I have been able to ascertain, all specimens of African
manatees in museums are from the rivers or coastal lagoons of the West
Coast between Senegal and the Quanza, and all of these specimens are
from regions below the first rapids. However, it is certain that such
specimens as are on record do not completely represent the distribution
of the manatees in Africa, just as it is certain that all published records
of their supposed distribution are unreliable. The waters from which
the manatees have been reported with certainty or probability of
correctness are the following:

SENEGAL

The African manatee does not seem to occur north of the Senegal
coast. Its presence there is established by the following records:

1757.—Adanson (p. 143) mentions the capture of manateesin the marigot (lagoon)
de Kantai in December and January.

1765.—Buffon (p. 390) notes the skull of a young specimen given by Adanson.

1793.—Pennant (II, p. 296) records the presence of a specimen in the Leverian
Museum.!

1836.—A specimen was collected by Robert (p. 363) for the Paris Museum.

1883.—Rochebrune (p. 190) states that the manatee was found in the marigots of
Lampsar, Leybar, and Bafing, and that Adanson mentioned their occurrence in the
marigot of Sorres, from which they have since disappeared. Rochebrune further
states that the manatee of the Senegal coast is not found in the rivers but in the
marigots.

1886.—Hartlaub (p. 15) records the presence of a manatee from Senegal in the
Vienna Museum.

1The contents of the Leverian Museum were disposed of by ‘auction : in 1806. A copy of the sale
catalogue with the buyers’ names added to some of the 7878 items is in the British Museum (Natural
History). An examination of this catalogue might reveal the present whereabouts of the specimen
named.

1934] Hatt, African Manatees 200

GAMBIA

Listed by Trouessart. Mr. Robert Rockwell informs me that while a
member of the Blossom Expedition he heard definite reports of the occur-
rence of manatees in the Gambia River.

SIERRA LEONE

1737.—Atkins (p. 42) notes the manatee in the Sierra Leone River.
1846?— Clarke (p. 128) lists “fishes”? commonly taken in Sierra Leone and among
‘them mentions the manatee.
1881.—Flower (p. 454) quotes the journal of one R. B. Dobree who was shown
places in which the manatees CAME ASHORE between Kikonkeh and the sea. Though
such a statement cannot be construed as good evidence for the occurrence of manatees
in the rivers of Sierra Leone, it seems to be probable that these animals do occur there.

LIBERIA

1885.—Biittikofer (pp. 144-147) recounts the capture of a manatee in Fisher-
man’s Lake behind Cape Mount, and states that this animal is found in Liberian
rivers up to the rapids.

1890.—Bittikofer (II, pp. 392-393) further records the occurrence of the manatee
in the vicinity of Millsburg, below the last cataract of the Saint Paul River, and adds
that in 1887 two were shot in the Missunado River.

1892.—Jentink (p. 199) records the presence in the Museum des Pays-Bas of
Biittikofer’s Cape Mount specimen.

Ivory Coast
CaRACALLA River: 1848.—Perkins (pp. 198-199) records a manatee from the

Caracalla River, 20 miles east of Cape Palmas. This specimen, the type of M.
nasutus Wyman, is now in the Museum of Comparative Zodlogy.

SLAVE COAST
1893.—The manatee is listed by Matschie (p. 180) in his ‘Mammals of Togoland.’
1897.—Thomas and Lydekker (p. 596) record specimens in the British Museum
from Lagos and Benin.

Nicer River SYSTEM

Manatees are known from three parts of the Niger system: the
lower reaches near the coast, the upper section above Timbuctu, and
from the river Benue.

Lower River: 1728.—Labat (p. 337) records that the ‘“‘manaty” was often
taken in the Niger. His is perhaps the first picture of an African manatee.

1857.—Baikie (p. 68) collected a specimen from the mouth of the Niger, and
Maclaud (1908, p. 289) writes that there is a Niger River specimen in the Paris Mu-
seum, though the section of the river is not stated.

Upper Nicer River: Though there appears to be no record of a manatee ever
brought out from the upper Niger River where it is bordered by great, marshy lagoons,
several reports make it seem almost certain that the animal is found in that region.

506 Bulletin American Museum of Natural History [Vol. LXVI

1858.—Barth (II, p. 605), who traveled along the upper Niger River, states that
the manatee occurs in the Isa near Timbuctu and (V, p. 103), in reference to the
decorations of some natives of the Niger above Timbuctu, he writes: ‘‘They wore also
a rich profusion of white rings which are made of the bones of that very remarkable
animal the ‘ayu’ (Manatus), which seems to be not less frequent in the western than
in the eastern branch of the Niger,” and (p. 472) ‘The lake [Débu, upper river, above
Timbuctu] besides fish, contains numbers of that curious animal called ‘ayu’
(Manatus).”’

1901.—Gratiolet (p. 248), on the report of M. Carpeaux of the Colonial Troops,
states that the manatee occurs at Zinder and at Segou, above Timbuctu.

1908.—Maclaud (pp. 289-290) reproduces a photograph of a manatee in the
Niger which he credits to the Mission Desplagnes, an archaeological expedition which
studied the Timbuctu region. Of course, it is only an assumption that this photo-
graph was taken in the upper river. Maclaud writes that the manatee is not uncom-
mon in the LAKES along the Niger near Timbuctu, but that this animal has almost
disappeared in the large coastal rivers.

1906.—Johnston’s (p. 246) observation that it would be remarkable if the
manatee could pass the Niger rapids from Busa to Say, seems to be answered by these
several reports.

1925.—I am informed by M. Lucien Blancou that in a book by Captain Pivert
titled, ‘Mes Chasses en Afrique et en Extreme Orient,’ published at Paris in 1925,
there is a photograph of a manatee taken in the Niger at Gao or Ansongo.

BENUE River: 1856.—Vogel (Owen, 1856, pp. 345-346) has described, and Owen
has named a manatee, the type of which is now in the British USE, and which
Vogel took in the Benue.

1857.—Barth (II, p. 605) records that he heard stories of this animal along the
Benue but did not see the animal.

1924.—Migeod (p. 167) found manatees common in this river especially at
Numan, below Yola.

1931.—M. Lucien Blancou, of the French Colonial administration, informs me in
correspondence that at Léré he saw articles made of manatee skin and that he has
seen a photograph of a manatee taken in Lake Léré. :

GuLF OF GUINEA

Outp CauaBarR River: 1860.—McBain (p 150) described a skull from Old
Calabar which is now (Turner, 1912, p. 156) in the Anatomical Museum of the
University of Edinburgh.

BrETIcKA-BA-MALLALE: 1897.—Sj6stedt (p .45) records the presence of manatees
at this point on the coast near Cameroon Mountain.

CAMEROONS: 1877.—Peters (p. 485) writes of a specimen taken by the Buchholz
Expedition near Wuri on the Cameroon River, and states that the manatee is also
found near the mouth of the river at Doctor’s Cape, as well as in a larger stream near
Mungo.

1886.—Hartlaub (p. 15, et seq.) records a specimen in the Berlin Museum and
one in the Liibeck Museum from the Cameroons. In a letter quoted by Hartlaub,
Pechuél-Loesche states that the manatee is found in the Camercons.

1897.—Three embryos from the Cameroons are described by Kiikenthal (1897a).

1934] Hatt, African Manatees Oot

1909.—Passarge (p. 446) states that the manatee occurs in the mangrove regions
of the Cameroons.

1917.—In his book on the Cameroons, Calvert publishes (Pl. cuxxvi1) a good
photograph of a manatee (labeled “walrus’’) surrounded by a group of natives, one
of whom holds a harpoon.

Rio Muni: 1886.—Hartlaub quotes a statement of Pechuél-Loesche that the
manatee occurs in the Rio Muni.

GABUN River: 1861.—Du Chaillu (p. 367). M. oweni here described is probably
from the Gabun River.

1865.—Gray (p. 1383) notes the presence in the British Museum of four skeletons
purchased from Du Chaillu and presumably from the mouth of the Gabun.

1886.—Hartlaub (p. 15 et seq.) lists three skulls from the Gabun River, one each
in Stuttgart, Hamburg, and Bremen museums.

OaGowE River: 1861.—Du Chaillu (p. 367) lists this as one of the rivers in which
manatees occur, and some of his specimens may be from here.

1886.—Hartlaub (p. 15 et seq.) notes four skulls from this river, now in the
Berlin Museum (Nos. 26333, 26335, 26337, 26338) and further states that Pechuél-
Loesche writes of their occurrence in the same stream.

Kouiuvu River: 1886.—Hartlaub (p. 99) quotes a communication from Pechuél-
Loesche to the effect that the manatee occurs in this river. However, in Pechuél-
Loesche’s account of the Loango Expedition (1882, p. 222), he states that no specimen
was taken in the region, though one was distinctly seen. Whether or not Pechuél-
Loesche received other records between the years 1882 and 1886, I cannot ascertain.

LuEmME River: 1889.—Noack (p. 105) quotes a Mr. Hesse to the effect that he
had a manatee-hide whip made from a specimen taken in the Luemme near Massabi.

CHILOANGA (TSCHILOANGA OR LOANGA) RivER: 1886.—Hartlaub (p. 99) quotes
Pechéul-Loesche as writing that manatees are found in the Chiloanga.

BELGIAN CoNnGco

Coneo River: 1746.—According to Barbot (p. 518) Merolla says that the Zair
(Congo) has plenty of these ‘‘monstrous fishes or mermaids, resembling a woman
upwards.”

1884.—Johnson (p. 379) writes that so far as is known the manatee never passes
the cataracts ot the Congo.

1886.—Hartlaub (p. 99) quotes Pechuél-Loesche to the effect that manatees occur
in the Congo.

1889.—Noack (p. 105) writes of the statement of a Mr. Hesse that the manatee
lives in the lower Congo, and quotes a seventeenth century account of manatees being
frequently taken in the Congo.

1890.—Bocage (pp. 29-30) writes of the manatee’s occurrence in the Zaire
(Congo) and states that the Lisbon Museum has a specimen from Angola, which may,
or may not, refer to the south bank of the mouth of the Congo.

1926.—Derscheid has given a valuable account of living specimens of manatees
from the Congo and has accompanied his notes with a good photograph showing one
of these out of the water resting on a platform in its aquarium. Dr. Rodhain is quoted
to the effect that the manatees are found in the Congo from Binda to the “‘Chaudron
d’Enfer.”’ It is also stated that the Musée du Congo Belge possesses a series of skins

008 Bulletin American Museum of Natural History [Vol. LXVI

and skeletons from this region. Scme observations on the natives’ hunting and
use of the species are given.

1930.—Schouteden (p. 370) states that the manatee is found in the lower Congo
only and that it seems to be particularly localized in the region about Boma.

The single specimen of a manatee collected by the Congo Expedition
was taken by Mr. Lang at Banana, in August, 1915. Photographs of
this animal are reproduced here as plate X XVII.

ANGOLA

1746.—Barbot (pp. 517-518) describes the manatee from the lakes of “ Angola,
Quihite, and Angolm in the province of Massingam,”’ localities not appearing on any
maps that I have consulted.

1920.— Possibly on information obtained from sources here quoted, Marquardsen
(p. 69) writes of the occurrence of manatees in Angola rivers.

Lose River: 1875.—Monteiro (p. 17) had part of a specimen from near Ambriz
at the mouth of the Loje.

Danve River: 1875.—Monteiro (p. 17) described the native method of trapping
the manatee near the mouth of the Dande.

BENGo River: 1897.—Greévé (p. 56) states that the manatee occurs in the Bengo,
but on whose authority this records rests, I do not know.

QuaNzZA RivER: 1875.—In this river, Monteiro (p. 17) saw a cance with much of
the flesh of a manateeinit. As faras I have been able to ascertain, this is the southern-
most coastal record of the manatee’s occurrence and is probably correctly founded.

UNCERTAIN OR ERRONEOUS REPORTS OF YT. senegalensis
Tue Lake CHapD BASIN

1858.—Barth (III, p. 289) wrote that ‘‘there is also in the river Shari a very large
animal apparently identical with the ayt of the Benue and Niger—Manatus Vogelit.”’

1881.—Nachtigal (p. 670), however, saw nothing of the manatee in the Shari
region, though he is said to have looked for it.

1924.—Migeod (pp. 147-167) was unable to confirm the existence of manatees in
Lake Chad, even though he made inquiry at several different places. The published
records of the Boyd Alexander Expedition, the Mission Tilho, Foureau’s Mission
Saharienne, and Chevalier’s Mission Chari-Tchad do not mention the animal.

1925, 1928.—The best evidence that the manatee does actually occur in Lake
Chad and the Shari appears to be found in certain references which I have not seen
personally, but which have been generously transmitted to me by letter from M.
Jacques Pellegrin of the Museum National d’ Histoire Naturelle. In substance these
references (Monod, 1928 and Pécaud, 1925) state that in the Chad basin the manatee
is now rare, but as elsewhere, was formerly abundant, that Major General Pécaud has
himself seen the skin of a manatee in the region and that he was assured by his aides
that previously, perhaps about 1905, the animals were captured near Fort Lamy.

1866.—In discussing the reputed occurrence of the manatees in the Chad district,
Murray (p. 420) states that in his opinion this distribution is quite possible because
‘“‘the watershed between the Lake and the Sea is not a lofty range of mountains from
one side of which the rivers run into Lake Tschad, and from the other into the Niger,

1934] Hatt, African Manatees 509

but a flat, marshy tract of land, so nearly level, that it is almost an equal chance by
which way the waters willrunfromit. It is lhkea large peat-bog, or a gigantic sponge,
out of one side of which creeps the Arre and Shari, and out of the other the Benue.
The Hippopotamus goes easily from one to the other, and in the rains, when the
country becomes flooded, the natives go about in boats. It is hke an inundation, so
that the manatee could with ease come up from the Atlantic and find its way into
Lake Tschad.”

THr Upper ConGgo SYSTEM

1874.—Schweinturth (II, pp. 159-160) heard from the natives of the Kibali, a
branch of the Uele, stories concerning a “‘river sheep”’ which he believed could be
nothing but a manatee. However, Schweinfurth believed that the Uele flowed into
Lake Chad and may have been influenced in his conclusion by Barth’s report.

1912.—From the Ubangi River tribes, von Wiese, Hauptmann, and Kaiserwal-
dau (I, p. 274) heard stories of a river animal which they assigned to the manatee.

1920.—Schwarz (p. 857) states that Schweinfurth, Schubotz, and von Wiese
heard of the manatee in the Mbomu and Uele. The reference to Schubotz, however, is
obscure, for I have found no mention of the manatee in his writings.

1887.—In his map of the distribution of the Sirenia (map 53f), Marshall accepted
the account of Schweinfurth and drew the conclusion that manatees occupied the
whole of the Uele and Congo.

1932.—M. Lucien Blancou states that the natives affirm the existence of the
manatee in the lagoons along the Likouala aux Herbes and in the Sangha, right bank
tributaries to the Congo in the region of Lukolela. Resting as this does on native
accounts their presence in these streams must remain, for the present, problematical.

Lakes Victoria, ALBERT, AND TANGANYIKA

1887.—Marshall (map 53f) indicates that Sirenia occur in the above-named lakes,
but on whose authority he indicated this fabulous distribution I do not know.

THe East Coast or AFRICA

Some authors of the nineteenth and twentieth centuries have stated
that manatees occur on the east coast from the Cape of Good Hope to the
Mozambique Channel. This must be due to credence given to reports of
early navigators who saw the dugong on the Madagascar coast or else-
where. There is on the Cape of Good Hope, between Cape Town and
Port Elizabeth, a Zeekoe River, this name, however, appears to refer to
the hippopotamus and not to the manatee or the dugong, for Sparrman
(1785, I, p. 346) speaks of this river harboring ‘‘sea cows (hippopotamus
amphibius, Plate 1v).” The animal figured is clearly a hippopotamus,
and Sparrman’s account of hunting ‘‘sea cows”’ further shows that this
animal was no sirenian. It is, however, stated in a letter which Sparrman
wrote to a friend, and which was subsequently published (Sparrman,
1777, p. 40) that the author captured a ‘“‘manatee”’ alive, and to judge
by the route of this traveler, the capture must have been made in Cape

560 Bulletin American Museum of Natural History [Vol. LXVI

Province. The possibility that the word ‘‘manatee”’ is an error in trans-
lation before publication must not, however, be overlooked.

ABYSSINIA

Lake Tana: 1868.—Heuglin (pp. 247 and 289) states that in Lake Tana there
lived a manatee-like animal called by a name which means “‘sea calf.’”’ That the
author did not confuse the alleged mammal with the hippopotamus is clear from his
statement that the hippopotamus also lived in the lake. Very large catfishes are said
to inhabit this lake, and it may be that these are creatures that gave rise to Heuglin’s
report.

MareEB River: 1877.—Heuglin (loc. cit.) heard reports of an animal similar to,
or identical with the supposed manatee of Lake Tana, living under a different native
name in the Mareb and its tributaries, a part of the Nile drainage system. For this
animal he suggested (1877, II, p. 187) identity with Manatus Vogelii of the Benue
River. It is perhaps superfluous to observe that these purported Abyssinian manatees
can only be considered mythical.

OcEANIC ISLANDS

St. HELENA: A marine mammal called a “‘manati”’ was formerly not uncommon
at St. Helena up to the nineteenth century, the last specimen having been killed in
1810. Lydekker (1899, pp. 796-798) concluded that this was in all probability a
sirenian, but not identical with the African manatees. Unfortunately, since no speci-
men of this creature is known, nothing is to be gained by speculation as to its probable
relationships, but it seems most likely that the conclusion reached by Dampier,
during his visit to the island in 1691, is the correct one. Concerning this he wrote
(1906, I, p. 526), “‘I was also informed that they get manatee or sea cows here, which
seemed very strange to me. Therefore enquiring more strictly into the matter, I
found the Santa Hellena Manatee to be by their shapes and manner of lying ashore on
the Rocks, those Creatures called Sea-lyons.”’

SUMMARY
There are three known living species of manatees: one African
(Trichechus senegalensis Link); one chiefly inhabiting the coastal waters
of the West Indies and the eastern American coast from North Carolina
to Brazil (Trichechus manatus Linnaeus); and a third occurring chiefly
in the rivers of northeastern South America [Trichechus inunguis (Nat-
terer)|. It is probable that geographic extremes of these species are
racially distinct, and one such, 7. manatus latirostris (Harlan), of
Florida is here recognized, but the manatee of the mouth of the Congo
cannot, on the basis of present knowledge, be distinguished from speci-
mens of senegalensis taken in Senegal, as figured and described in the
literature. 7. senegalensis resembles T. manatus more than it does
inunguis, and it is supposed that these two species are more closely

related than the two New World species.

1934] Hatt, African Manatees 061

A study of the skeletons of the species of manatees shows that the
principal postcranial differences lie in the proportion of the elements of
the pectoral girdle. 7. senegalensis has lightly built bones and hands
proportionately about as long as those of manatus. ‘The pectoral limb
bones of manatus are about double the diameter of those of the African
manatees, though their proportionate length is similar. The limb bones
of inunguis are lightly built like those of senegalensis, but the distal
elements, particularly the metacarpals and first phalanges, are consider-
ably more elongate than those of the other two species.

The sterna of the three species are of distinctive types. ‘These are
shown in figure 1.

The differences in the skulls, for knowledge of which we are mainly
indebted to Hartlaub, are centered principally about the nasal basin.
Individual variation, which is so extensive in the genus, hinders the
formulation of invariable criteria for the distinction of the species, but
this variation is most pronounced in such vestigial parts as the lacrimal
and nasal bones. Perhaps the most constant specific character of the
skull is the length of the vomer. In senegalensis this usually extends
only to the level of the middle of the orbits, in inunguzs to the anterior
edge of the orbits, and in manatus to the posterior edge of the incisive
foramen or beyond.

Mandibular characters are well marked, though, in the main, subject
to considerable age variation. The species znungwis is easy to recognize,
among other features, by the large number of mental foramina (ten or
more), manatus by its deep symphysial groove, and senegalensis by its
lack of such a groove.

The range of the African manatees extends along the West African
coast from Senegal to the Quanza River in Angola, and this full coastal
range is represented by scattered specimens in the museums of Europe.
The manatees are not known from far up any of the rivers flowing into
this coast, if we except the Niger and its tributaries. A specimen in the
British Museum was found along the Benue, and published reports
make it seem certain that manatees occur in the upper Niger River
above Timbuctu. It is also probable that manatees have occasionally
been taken in the Lake Chad drainage, into which they are supposed to
have gained access by crossing the seasonally flooded area lying between
the Benue and the Shari. Reports of manatees in the Uele, Lake Tana,
east of the Cape of Good Hope, and St. Helena are discredited.

5962 Bulletin American Museum of Natural History [Vol. LXVI

BIBLIOGRAPHY

Apanson, M. 1757. ‘Histoire Naturelle du Sénégal.’ 190 pages. Paris.

ATKINS, JOHN. 1737. ‘A voyage to Guinea, Brasil, and the West Indies.’ 2d Ed.,
xxv +265 pages. London.

BalkIE, BaLtFour. 1857. ‘On the skull of a Manatus from western Africa.’ Proc.
Zool. Soc. London, pp. 29-33.

Barsot, J. 1746. ‘A description of the coasts of north and south Guinea and of
Ethiopia Inferior . . .,’ in Churchill and Churchill’s ‘ A collection
of voyages and travels.’ V, 708 pages. London.

BartH, Hernricw. 1857-1858. ‘Reisen und Entdeckungen in Nord-und Central-
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564 Bulletin American Museum of Natural History [Vol. LXVI

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1934] Hatt, African Manatees 565

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ROBERT,

066 Bulletin American Museum of Natural History [Vol. LXVI

ADDENDA REGARDING LAKE CHAD MANATEES

Since the date of completion of the manuscript (March, 1932) of
this bulletin there has been published a photograph of a manatee (1933,
Bulletin Economique de |’Afrique Equatoriale Frangaise, 9° Année,
No. 30, p. 35) with the caption ‘“‘Un lamentin péché dans le Tchad.”
Inquiry has elicited the information that this photograph was taken
in the Chad district, but not in the Chad drainage. It was in fact
secured in the lake at Léré, in the vicinity of Moyo—Kebbi, from which
records have already been quoted.

Puate XXVIII
Figs. 1 to 3. Manatee at Banana, Belgian Congo, A.M.N.H. No. 53989.

Buttetin A. M. N. H. Vou. LXVI, Prats XXVII

— Tun PANGOLINS aND AARD-VARKS
— COLLECTED BY THE AMERICAN
_ Museum Congo EAESOTTION

Co?
Co
ua el

2 Be ae tT -HATT =~

BULLETIN :
_OF
“THE AMERICAN MUSEUM OF
NATURAL HISTORY

- VOLUME LXVL. 1934
_ ARTICLE VII :

NEW YORK
“September 14, 1934

59.9,31 M (67.5)
Article VII—THE PANGOLINS AND AARD-VARKS
COLLECTED BY THE AMERICAN MUSEUM
CONGO EXPEDITION!

By Rospert T. Harr

Puates XXXII to XXXIX; TEext Figures 1 anp 2

CONTENTS

PAGE
“OPTED TOG TEIO INS 5 Hd iltbace: ees a Seat RN Se ISLET cet re en a a 643

Species with Their Localities and Number of Specimens from Each
OG AI Teen ce) ee eine ai due arate a Sh OS Aa rye 645

List of Localities, with Names of the Species and Subspecies and Number
Smspeemmensmlaken atiwMach Locality... 23. 3. 2029522352450 ee 645
Newssubspecies and Its Type Locality... . 02... 2... ese ce ee eee 646
HON SEETTEUST. oo oc oe tS CRIS ee OO eters a SS oe 646
ASICEDES CHARCOAL So's boc, 3 ce Se acta ae DEA aa ee EE re i ier nee 646
PDH TSMLONOUCHUGAUUSRO er acti tra, onus aARee tt: aah * oS Leia Wisc mnt “OO
Manis tricuspis. . eee ee ED
Distinctions between ihe Skulls of M. Ueelepis and M. Luaicaudatis. 659
ORYCTEROPODIDAE. . oa: SN Ss Copan Ned ee cen ha eT PRIN 6 |
Or icrpus SPRSaOt Peon. eT RTE Le ONO RL Fra rat ad Anh S On 662
OHAGLERO DUS: CLUKSSONM/ONADIUUSS nse veo yo cee no SONS ee een ee 663
ECVE LT OGTR UNDISR Toe 5 5 alle cee py dhe ORE OL Bet ou cn I ea Re a ea a a 671

INTRODUCTION

The collections of pangolins and aard-varks made by Messrs.
Herbert Lang and James P. Chapin, while functioning as the American
Museum Congo Expedition, are of exceptional value in that the spe-
cies represented were secured in sufficient numbers to enable one,
apparently for the first time, to gauge the range of variation occurring
within geographically limited populations of these species.

There have been many revisions of the Manidae between that of
Sundevall in 1843 and that of Frechkop in 1931, and current nomencla-
ture synthesizes their conclusions. Thus, the two African arboreal
species and the giant pangolin are now commonly recognized as generical-
ly differentiated and without subspecies, though exhibiting great indi-
vidual variation in some characters, a status which the ninety-five scaly
anteaters of the collection does not affect.

The Orycteropodidae, to the contrary, appear to present noteworthy
localization of character combinations within areas not far separated.

iScientific Results of the Congo Expedition. Mammalogy, No. 15.
643

644 Bulletin American Museum of Natural History [Vol. LX VI

To the long list of names for this latter animal already carried in the
literature, it was deemed advisable to add another, though it is realized
that our knowledge of the speciation of Orycteropus is totally inadequate.
Unfortunately the meager material in the museums of this country does
not place me in a position to judge as to the validity of the many forms
described, often insufficiently or on the basis of a single specimen.

All of the “species” and “‘genera”’ of African pangolins named by
the early workers, which are, in my present opinion, without foundation,
have been placed in synonymy by reviewers of the Order, and for this
reason I have not burdened this paper with a detailed synonymy. My
one departure from currently accepted ranking of names of the scaly
anteaters is the relegation of the names Phataginus, Smutsia, and Uro-
manis, recognized by Pocock and others as full genera, into subgeneric
position. I do this with no depreciation of the established fact that the
groups in question are very distinct, nor without appreciation of the
importance of the characters upon which the distinction is based. It
does seem, however, that nomenclature gains nothing and loses much
by the multiplicity of generic names where each genus of a single family
is represented by a single, or at the most, only two species. I consider it
advisable to recognize these differences by name, but not so to elevate
the rank involved that the fundamental relationship of all forms is
obscured by designations that can be of meaning to no one but the
specialist.

In the Congo Expedition report on ants there was published a sec-
tion by Dr. J. Bequaert on ‘The Predaceous Enemies of Ants,’ to which
Mr. Lang contributed a full and interesting section on mammals. This
was in part founded on his field notes, but these were not completely
given and not always recognizable as original. For this reason it seemed
appropriate to reproduce them here, without additions, for their value as
original information. In certain instances, as indicated by quotation
marks, these notes are directly transcribed. Some notes, however,
have been rewritten in order to place the subject matter in a more
convenient form.

The excellent photographs of living and recently living animals
(Plates XXXII to XX XIX) are products of Mr. Lang’s painstaking
photography. His labors resulted in many pictures of these species, of
which those reproduced are but a small selection. Text figures 1 and 2
are from the pen of Miss Margaret M. Matthew.

It is a pleasure to acknowledge the generosity of Dr. Glover M.
Allen of the Museum of Comparative Zoélogy, and Dr. Gerrit 8. Miller,

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 645

Jr., of the United States National Museum, who loaned me such speci-
mens as were in the collections in their charge. The manuscript of this
report has received a critical reading by Mr. H. E. Anthony, and I wish
to express my appreciation of his numerous helpful suggestions.

SPECIES WITH THEIR LOCALITIES AND NUMBER OF SPECIMENS FROM HACH

LOcALITY
SPECIES LOCALITIES SPECIMENS
1. Manis (Smutsia) gigantea Bafuka, 1; Niangara, 1; Niapu,
12; Poko, 2. 16
2. Manis (Uromanis) longicauda- Akenge, 2; Bolobo, 1; Gamangui,
tus 2; Medje, 1; Niapu, 7. 13
3. Manis (Phataginus) tricuspis Akenge, 15; Avakubi, 6; Faradje,

1; Gamangui, 1; Medje, 20;
Ngayu,1; Niangara, 2; Niapu,

18; Poko, 1; Stanleyville, 1. 66
4. Orycteropus ertkssoni ertkssona Niapu, 2. 2
5. Orycteropus ertkssoni faradjius Faradje, 11. 13

List oF LOCALITIES, WITH NAMES OF THE SPECIES AND SUBSPECIES AND
NUMBER OF SPECIMENS TAKEN AT EAcH LOCALITY

No. oF
LOCALITIES SPECIES AND SUBSPECIES SPECIMENS TOTALS
Akenge Manis longicaudatus 2
" Manis tricuspis 15 17
Avakubi Manis tricuspis 6 6
Bafuka Manis gigantea 1 IL
Bolobo Manis longicaudatus 1 1
Faradje Manis tricuspis 1
Orycteropus e. faradjius il 12
Gamangul Manis longicaudatus a
os Manis tricuspis 1 8
Medje Manis longicaudatus 1
i Manis tricuspis 20 21
Ngayu Manis tricuspis 1 ]
Niangara Manis gigantea 1
* Manis tricuspis 2 3
Niapu Manis gigantea 12
5) Manis longicaudatus 7
Mamas tricuspis 18
s Orycteropus e. erikssona 2 39
Poko Manis gigantea 2
- Manis tricuspis 1 3
Stanleyville Manis tricuspis i 1

646 Bulletin American Museum of Natural History [Vol. LXVI

New! SuBsPeciIgES AND Its Typr LocALity

1. Orycteropus ertkssoni faradjius Harr. Faradje.

MANIDAE
Manis Linnaeus?

Manis LinnaEvs, 1758, ‘Systema Naturae,’ 10th Ed., p. 36. Type by mono-
typy, pentadactyla.

The following names, which, among others, have been proposed for
the African pangolins, are here recognized as subgenera.

Phataginus RaFINESQUE, 1821, Ann. Gén. Sci. Phys. Brux., VII, pp. 214-215.3
Included species tricuspis and ceonyx (=longicaudatus). Type by subsequent designa-
tion (Pocock, 1924, p. 722), tricuspis.

Smutsia GRAY, 1865, Proc. Zool. Soc. London, 1865, p. 369. Type by designa-
tion, temminckit.

Uromanis Pocock, 1924, Proc. Zool. Soc. London, p. 722. Type by designation,
longicaudata.

Manis (Smutsia) gigantea I]liger
Plates XXXII to XXXIV

Manis gigantea IuuicER, 1815, Abh. d.k.Ak.d. Wissensch. Berlin, p. 84.

?Pholidotus wagnert FirzInGER, 1872, Sitzber. K. Akad. Wien Math. Naturwiss.
Cl., LXV, (1), p. 48.

Manis gigantea is a terrestrial pangolin measuring in excess of 1200
mm. when adult. In common with all other African pangolins no hairs
project between the scales, the median dorsal row of scales does not
extend to the tail tip, and there is no external pinna of the ear. The
belly is naked, the preaxial surface of the fore limb bears scales to the
base of the claws, the tail is massive and bears no naked subterminal
pad, characters shared with a smaller species temminckii. There are
12 to 15 scales in the median dorsal row of the tail. The massive skull
measures over 130 mm. in adults.

Represented by 14 skins, 11 skeletons, 1 skull, and two fetuses in
formalin, collected as follows:

Bafuka, 1 (skin of adult), April 5, 1913

Niangara 1 (adult 2), April 26, 1913

Niapu, 12 (6 adult <7, 4 adult 2, 2 fetuses), November 18, 1913-
January 27, 1914

Poko, 2 (1 adult @, 1 im. 9 ) August 4 and 10, 1913.

iHatt, Robert T. 1932. ‘The Aard-vark of the Haut Uele.’ Amer. Museum Novitates, No. 535, p. 1.
2The reader is referred to Pocock (1924, p. 718) for a review of the generic nomenclature of the

pangolins.
3The title-page bears the date 1820, but according to Sherborn (see Pocock, 1924, p. 721) it was not

published until the following year.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 647

The accumulated observations of the past century show profound
differences between gigantea and its nearest relative, temmincki, but
due either to a remarkable uniformity of the species or to the rarity of
specimens, no one has yet pointed out any geographic differentiation
within either species. The Congo Expedition series, representing the
eastern limit of the range of gigantea, forms a basis for the judgment of
range of variation found in specimens from other localities.

Sizm.—The external measurements—average (minimum-maxi-
mum)—of twelve adult Manis gigantea, taken from animals in the flesh,
are as follows:

TotTaL LENGTH Tart LENGTH
1 ee, © 1373 (1185-1530) 627 (550-700)
5 ot 1438 (1370-1530) 674 (650-700)
7 © 1298 (1185-1365) 596 (545-675)

The maximum size of the species appears not to be represented by
this series, for Schubotz (1912, p. 356) has recorded a specimen 1650
mm. long from Angu, and Thilonius (1912, p. 373) mentions one 1600 mm.
in length. Also there is a skull from Liberia in the collections of the Mu-
seum of Comparative Zodlogy that is slightly larger than any in the
Congo collection.

Cotor.—The color and color pattern show some divergence in the
series that is in part traceable to age changes but which is also somewhat
the result of differences in methods of preservation. The greater
number of the skins have been tanned and are thus free of foreign color-
ing matter, but a few have not received further treatment than that
which was given them in the field, and, in consequence, the furrows on
the bases of the scales harbor a considerable amount of the red soil of
the region, and the general color of the animal is altered.

In the adults of medium size a color pattern is attained which is the
same in both sexes and may be considered typical of the species. In a
specimen of such an animal the scales of the head, neck, shoulders, arm,
and hind legs are dominantly dark olive-brown.! This color shades
gradually into avellaneous over the dorsal region. Individually the
head scales are uniformly heavily pigmented except for light marginal
bands of buffy brown. The dark area is more and more restricted toward
the rear, so that in the mid-dorsal region the deep olive is confined to the
basal and medial superior surface. This is flanked by a variable brown-
ish area that is near buckthorn. Over the tail a deep Roman green
assumes increasing prominence in the apical part of the scale. Here the
base of the scale is a light brown.

1The color terms refer to Ridgway’s ‘Color Standards and Color Nomenclature.’

648 Bulletin American Museum of Natural History [Vol. LX VI

Though individual variation and differences in method of preserva-
tion seem to account for a high percentage of the differences represented,
there is some divergence that is obviously due to age and its attendant
wear on the scale surface. This attains extremes in the young animals
with rough, unworn scales and in the old individuals in which all seales
are smooth and polished through friction between themselves and be-
tween the scales and their environment. In both of these age extremes
the color, general and local, is lighter. In the young animals this is
achieved through the masking occasioned by rugosity and soiling, and
in the older individuals, it seems, through the wearing away of the pig-
mented layer, though this of course may be only one of the factors
responsible.

The naked skin of the under parts is, in the tanned hides, near warm
buff. In his field catalogue Mr. Lang described a freshly killed animal
in the following terms: ‘‘Nose dull blue-gray, rest of snout and other
naked parts on head, pinkish gray, ventral surface grayish white. The
iris darkish.”’

ScaLeE TopocrapHy.—It has been written (1931, Frechkop, p. 7)
that the scales of gigantea are without keels, but this is not wholly true,
for the two smaller specimens of gigantea in the Congo series, where wear
of the scales is very limited, show well-developed and characteristic
median keels on the scales of the flanks and the limbs (Plate XXXII,
figure 1). These are also prominent in the same areas in the embryonic
specimens of the species (Plate XX XIV, figure 1). Keels are also found
on the lower flank scales of young specimens of temminckiz, however,
and it would thus seem probable that these are characteristic of all of
the scaly anteaters, and that the early disappearance in the large species
is due only to the greater wear to which these terrestrial forms are
subjected.

The shape of the scales in the two species of the Smutsza group 1s
altered by the wearing away of the tip. In the dorsal region the pos-
terior scale border is occasionally perfectly transverse, and the scales of
both the back and tail appear as though truncated by some artificial
process. More commonly the free margin is modified into an elipsoidal
are.

Scate NumMBER.—Scalation is relatively constant, but not exactly
so. The full number of scales appears during uterine life, and modifica-
tion in scale count is, it seems, occasioned only by injury. Several
individuals which were obviously confined for some time by a devise in
the form of a halter have lost the scales in lines where the restraining

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 649

cords cut in. One old male from Niapu (53854) possesses a large number
of small accessory scales largely underlying, though in some instances
projecting beyond, a normal scale. These supernumerary scales vary
from styliform to falciform. None approach the normal form or size for
the region in which they are located.

The scale counts of gigantea and temminckii have been reported by
others, and the present series adds little to our knowledge concerning the
variation in these numbers.

Hair.—The species is hairless, except for a dense ring of short,
circumorbital bristles and a patch of similar hairs in front of the audi-
tory meatus. No other trace of hair is found on either the dried skins or
the embryos preserved in alcohol.

SKULL.—Within the series of gigantea skulls from the Congo collec-
tion there is little variation due to factors other than age, sex, and injury.
One female (53853) presents great asymmetry in the occipital region, and
in another (53851) there is extensive malformation in the nasals and
frontals due, apparently, to injury occasioned by the long-continued
restraint of a tightly binding halter. The second and third cervical
vertebrae of this specimen are fused, the result certainly of abnormal
conditions. Two of twelve skulls show bregmatic bones. The dorsal
profile is in some crania very nearly straight, but in others it presents a
marked depression in the frontal region. (Contrast Nos. 53849 and
53846.) This seems, however, not to be correlated with age or sex.
Among the features presenting extensive individual variation that are
seen in the basal surface of the skulls are the configuration of the ventral
margin of the foramen magnum, the shape and size of the interpterygoid
fossa, the shape of the palatine notch, the caudal extension of the pala-
tine processes of the maxillae, and the presence of a palatine surface on
the vomer.

The only sexual difference which I have found in the skulls is the
attainment of greater size in the males and the sutures closing in the
females when the skull is smaller than at the time corresponding closures
occur in the males.

Age changes involve little other than increase in size, thickness and
density of bone, closure of sutures, and reduction in the height of the
lateral palatine ridge. The closure of the principal sutures in and across
the median plane is approximately as follows: dorsomedian suture of
exoccipitalia; basioccipital-basisphenoidal and the basisphenoidal-
presphenoidal suture; frontal suture; nasal suture (occurring in old
age).

650 Bulletin American Museum of Natural History [Vol. LXVI

Cranial Measurements of Adult Manis gigantea

Average (minimum-maximum)

4o 6 9 Oeil; ©

Greatest length 152 (148-162) 142 (184-148) 146 (134-162)
Basal length 142.7 (188-155) 183 (127-139) 137 (127-155)
Palatal length 94.5 ( 89-105) 90 ( 86-99) 92.4 ( 86-105)
Breadth across zygomatic

processes 49.5 ( 49- 50) 47 (45-49) 48.3 ( 45— 50)
Greatest breadth of braincase 55.6 ( 54- 57) 53.4 ( 51— 56) 54.4 ( 51— 57)
Greatest length of nasals 59.2 ( 56— 62) 52.8 ( 51— 61) 55.6 C5I- 62)
Breadth of single nasal 13 (12-14). 12:4 ( 12 13), aes)

Greatest length of mandible 112.5 (109-121) 105.5 (100-110) 109.38 (100-121)

Digestive TrRact.—When in the field Mr. Lang observed that in
a female 1360 mm. long that ‘‘The intestine measured 1080 mm. There
is no caecum. The stomach is practically divided in two sections: one
section is surrounded by strong muscles and resembles a gizzard; it
was filled with small stones (the largest 5 mm. in diameter) and heads of
ants it had eaten. There is towards the middle a large, roundish, raised
gland patch; the rest of the stomach is provided with folds; and has
been preserved in formalin. Towards the pyloric end is a raised portion
that looks like a sort of stopper, and as it projects far into the stomach it
certainly is of some assistance in grinding the food along its roundish
folded surface.”

ANAL GLANDs.—“‘ On either side of the anus are two ducts from which
a strongly smelling, white pasty excretion is given off on pressure from
the outside,” according to Mr. Lang.

REPRODUCTION.—Two fetuses were found in specimens of gigantea
measuring 1350 and 1365 mm. in total length. One of these fetuses
taken November 28, measures 240 mm. in total length. The other (Plate
XXXIV, figure 1) removed on December 9, measures 290 mm.

ScaLes, Drerense.—‘‘The scales on the sides hang very loosely
down when [the animal] stretched out,” wrote Mr. Lang. ‘If one tried
to unfold the living specimen it would suddenly glide its tail along its
side, often with such force as easily to bend over a native-made knife of
3 mm. thickness stabbed in its side.”

Burrow.—A burrow of this species, photographed by Mr. Lang at
Babeyru, is shown in Plate XX XIV, figure 2.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 651

Manis (Uromanis) longicaudatus (Brisson)
Plate XX XV; Figure 1

Pholidotus longicaudatus Brisson, 1762, ‘Regnum Animale,’ p. 19.

Manis tetradactyla LINNAEUS, 1766, ‘Systema Naturae,’ 12th Ed., p. 53.

Manis macroura ERXLEBEN, 1777, ‘Systema Regni Animalis,’ p. 101.

Manis africana DESMAREST, 1820, ‘Mammalogie,’ part 1, p. 376.

? Manis ceonyx RAFINESQUE, 1821, Ann. Gén. Sci. Phys. Brux., VII, p. 215.

Manis longicauda SUNDEVALL, 1843, K. Vet. Akad. Handl., 1842, p. 251.

Manis guineensis FiTzINGER, 1872, Sitzber. K. Akad. Wien Math. Naturwiss.
Cl., LXV, p. 24.

Manis senegalensis F1TzINGER, 1872, idem, p. 25.

Manis hessi Noack, 1889, Zool. Jahrb. Abt. Syst., IV, p. 100.

Manis longicaudatus is an arboreal species with a long prehensile
tail, equaling about two-thirds of the total length. There is a naked
area on the under surface of the tail tip. The forearms bear no scales,
but are covered with hair. The scales are large, yellow, and on the
flanks are keeled. The two inferior postscapular scales are markedly
larger than those adjacent to them. The skull may be recognized by its
sharp anteorbital narrowing.

Represented by 10 skins, 3 skeletons, and 8 skulls, collected as
follows:

Akenge, 1 (adult o) October 10, 1913

Bolobo, 1 native skin, December, 1914

Gamangul, 2 (adult o and adult 2) February 3 and 13, 1910

Medje, 1 (juv.) June 9, 1910

CoLLEcTOR’s MEASUREMENTS

Tota LENGTH Tait LENGTH Foot LENGTH
Io, 2 860.4 (755-937) 577.0 (505-645) 50.4 ( 44-55)
3 0 874.0 (810-937) 613.3 (560-645) 53.0 ( 51-55)
5 Q 851.4 (755-930) 560.6 (505-623) 49.0 ( 44-54)

Other measurements made by the collector on an adult male of 810
mm. total length, collected at Gamangui, are:

Middle of eye to tip of snout 35 mm.
Middle of eye to ear 25 mm.
Transverse diameter of mouth 18 mm.

EXTERNAL CHARAcTERS.—There is a low coefficient of variability
in scale count and color of longicaudatus. All of the specimens lie within
the range of scale numbers set forth by Frechkop. In color the only
noteworthy deviation from uniformity is the sporadic occurrence of a

652 Bulletin American Museum of Natural History [Vol. LXVI

few blond scales, usually on the tail but occasionally occurring on the
flanks. That this lack of basal pigmentation in these few scales is not
due to post-mortem desiccation or other changes is demonstrated by
their visibility in a photograph of a specimen in the flesh (Plate XX XV,
figure 2). Preserved skins often show a pronounced difference in color,
due to soiling, that disappears when the scales are wetted.

The belly hair is black in most individuals, but in some is bleached or
entirely changed to tawny or to russet. In none of the specimens ex-
amined does the throat or chin hair show any deviation from black.
The hair over the dorsum of the antibrachium varies from black to russet,
the usual combination being a black center stripe flanked by russet.

The external characters of a fresh specimen obtained by Mr. Lang
were described in his field notes as follows: ‘‘The whole face, inclusive of
nose and ears, dark brown, nearly black. The eyelids are thick and
protruding, the eyes themselves are small and appear brilliant black.
The toenails are blackish, the pads brown. The naked skin on the under
side near fore limbs pinkish brown. The naked skin about the anal
region pinkish gray. The scales look as nice and clean as if they had been
rubbed off with oil. The anal region is shghtly protruding, the penis
hardly visible, the testes underlying the skin are imbedded in the fatty
tissue outside of the abdomen (without forming a scrotum).”’

AGE CHanGcE.—In this species the alteration of the scales with
advancing age does not follow the same pattern as that of the other two
pangolins secured. The scales of the back retain the regular contours of
youth until extreme old age, when, to judge principally from a large
male specimen in the United States National Museum (No. 220402),
collected by Mr. C. R. Aschemeier near Fernand Vaz, the scales are
arrested in their growth and become scarred, broken, and irregular. The
small scales of the head, feet, and under surface of the tail of this speci-
men are so extremely worn that in many cases they are reduced to
little polished nodules presenting no free edge.

There is a tendency for the keeled scales of the sides, crus, and under
tail to become mucronate and to parallel to a shght degree the tridentate
scale shape of tricuspis. This is most pronounced in mid-life.

Color changes of the scales are confined to the shoulder and head,
where a gradual darkening occurs at the base of these growing structures.

The hair over the belly of the older individuals is longer and denser
than that in younger individuals from the same area.

GEOGRAPHIC VARIATION.—Representatives of the long-tailed pango-
lins from areas other than the Medje forest district do not always fall

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 653

completely within the limits of variation exhibited by our series from
that region. Thus the specimen from Bolobo (a native skin) and one
collected at Lukolela by Mr. Franklin Edson, III, both have the middle
front claw 5 and 2 mm. longer, respectively, than any seen on specimens
from other districts. The hair on the chins of these two animals from
the Moyen Congo is also longer than that of the specimens from the
forest of the Ituri-Haut Uele border, but in the absence of associated
variables, and with such small series to study, there is no justification

pu”
TT ila A.M.N.H.53 861

Fig. 1. Extremes of skull shape in Manis longicaudatus. a.—Immature 9°,
Akenge, Belgian Congo (No. 53861); b.—Immature ¢, Gamangui, Belgian Congo
(No. 53864).

for assigning distinctive names to the animals bearing these minor
characters. It may be added that the few specimens from Liberia, the
Cameroons, and Gabun that I have examined show no single feature
not duplicated by some specimen in the small series of the Congo collec-
tion, if marks of advanced age, discussed above, are excepted.

THE Skuity.—The skulls of Manis longicaudatus vary to the ex-
tremes of proportion exhibited by two female specimens taken within
40 kilometers of each other, shown in figure 1. These two are of only

654 Bulletin American Museum of Natural History [Vol. LXVI

approximately the same age, the short, wide skull having come from an
individual 890 mm. long, the other from one 795 mm. The skulls them-
selves are within one millimeter of being the same in greatest length, but
in one (53861) the width is but 44 per cent of the length, while in the
other (53864) the width is 52 per cent of the length. That the disharmony
exhibited is not due to age differences is shown by the fact that the oldest
individuals in the complete series are nearer the shape of the skull from
the smaller animal than that from the larger, and that in this series there
is a perfect gradation from one type to the other among animals of very
nearly the same size. The difference is not one of sex, since these two
are females, and the other skulls available show no sex-form relation-
ship. That two species, or races, are not involved is attested by the
uniformity of other characters and the random distribution of the skull
types within the localities represented by the Congo collection and other
material coming from points ranging between Liberia and the middle
Congo. Neither do the specimens examined lend support to any assump-
tion that one type or other is the result of a pathologic condition or an
endocrine disturbance, unless disruption of normal development is
chronic in the species. Other matters of random variation in the skulls,
such as the presence of Wormian bones, the variability of position in
sutures, etc., appear to be of about the same magnitude as variation
found in the better series of tricusprs. It must, therefore, be concluded
that in development the skull of longzcaudatus is normally subject to a
wide variability.

The greatest age is not represented in the Congo series. The
United States National Museum specimen (No. 220402) from Fernand
Vaz, described above for old age characters of the scales, is accom-
panied by a skull 6 mm. longer in greatest, basal and palatal lengths,
2 mm. longer in nasal length, and 4 mm. longer in mandibular length
than the largest skull of the Congo series.

The fact that the two male skulls are longer than the seven female
skulls in the same collection is obviously due in part to the accident that
the young individuals were all females, but it is probable that in this
species, as in the two others considered here, the males do attain greater
size than the complementary sex.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 655

Cranial Measurements of Manis longicaudatus

Greatest length
Basal length
Palatal length

Breadth across zygomatic

processes

Greatest breadth of brain

case

Greatest length of nasals

70.0 (69.1-71.0)
65.6 (65.5-65.7)
40.6 (40.2-41.0)

24.0) (23.6—24.4)

32.8 (31.4-34.2)
25.0 (24.1-25.9)

Average (Minimum-maximum)

7 2
65.3 (62.1-68.7)
60.2 (57.0-64.4)
36.9 (34.6-39.0)

24.5 (22.8-26.9)

21.3 (28.5-33.7)

10 #, 2

65.8(62.1-71.0)
60.8 (56.0-65.7)
37.1 (32.0-41.0)

24.2 (22.0-26.9)

31.5 (28.5-34.2)

22.5 (19.1-25.8) 22.8 (19.1-25.9)
5.6( 5.4- 5.8) 5.6( 4.5- 7.5) 5.4 ( 4.2- 7.5)
42.6 (41.2-44.0) 44.0 (38.2-51.0) 43.3 (38.2-51.0)

Breadth of sing!e nasal
Greatest length of mandible

Manis (Phataginus) tricuspis Rafinesque
Plates XXXVI and XXXVII; Figure 1

Manis tricuspis RAFINESQUE, 1821, Ann. Gén. Sci. Phys. Brux., VII, p. 215.
Manis multiscutata Gray, 1843, Proc. Zool. Soe. London, p. 22.
Manis tridentata Fociu1.0n, 1850, Rev. de Zool., (2) II, p. 472.

Manis tricuspis is an arboreal species with a tail constituting over
half the total length. The characters of the tail tip and the fore limbs are
like those of Manis longicaudatus. ‘The scales, however, are small and
numerous, brown, and during mid-life, tricuspid. The postscapulars are
not enlarged. The skull is broad-muzzled, and its zygomatic processes
are large and divergent.

Represented by 59 skins, 12 skeletons, 49 skulls and 2 embryos in
alcohol, collected as follows:

Akenge, 15 (4adult o,9 adult 2,1 adult sex ?, 1 juv. 2), September
10—October 22, 1913.

Avakubi, 7 (8 adult o’, 3 adult 2, 1 embryo), October 29, 1909-
December 8, 1913.

Faradje, 1 (adult 2), March 24, 1912.

Gamangui, 1 (adult o), February 11, 1910.

Medje, 20 (9 adult o’, 7 adult @, 2 adult sex ?, 2 juv. &), January
24—October 2, 1910.

Negayu, 1 (adult o), December 19, 1909.

Niangara, 2 (adult o’, adult 2), December 2, 1910 and April 20,
1913.

Niapu, 18 (6 adult o’, 8 adult 9, 4 juv. 2), November 10, 1913-
January 3, 1914.

Poko, 1 (adult o'), August 11, 1913.

Stanleyville, 1 (adult o), August 9, 1909.

656 Bulletin American Museum of Natural History [Vol. LXVI

EXTERNAL CHARACTERS.—The external appearance of a living
adult female, taken at Avakubi, November 10, 1909, was described by
Mr. Lang in the following words: ‘“‘Snout and skin around eyes pinkish
brown, lips pinkish, iris dark brown. The skin that may be seen under-
neath the hair behind the eyes and around the ears is grayish. The
visible skin on the fore and hind limbs dirty gray; throat, abdomen, and
skin around anus, also the underside of legs, grayish white.” The
collector recorded the eye color of another female as medium brown and
noted that the pupil was circular. The nose of a third female is described
as ‘“‘erayish above, somewhat brownish.”’

A male taken at Stanleyville, August 9, 1909, was characterized as
follows: ‘“‘Snout pinkish gray, underlip whitish, iris dark brown, eyes
much protruding from the thick, swollen looking eyelids. Ears can be
closed though they are often open, a simple slit outside, but supported
by a cartilage.”

AGE CHANGES IN EXTERNAL CHARACTERS.—The splendid series of
tricuspis from a small area in the upper Congo basin shows extreme limits
of variation in the contour, size, and count of the scales and the color,
length and density of hair that embraces practically the entire range of
variation that I have seen in specimens from Liberia, the Ivory Coast,
Cameroons, Fernando Po, Gabun, the lower Congo, Kasai district, and
central Angola.

The scales do not increase in number but grow in length and breadth.
Throughout life they are deeply striate, but in old age the scales of the
head and the tips of all scales, become worn smooth. At birth, and from
that time until the individual attains a length of about 325 mm., the
margins of the scales are even, but with ensuing wear, which by action
between the scales is concentrated on the section bordering the median
keel, the scales become sharply dentate, or, usually later, tridentate.
This configuration of the scale margins is characteristic of youth, and
usually disappears, particularly dorsally, by the time the individual
reaches a length of 750 mm. During the period from youth to old age
the scales grow in length more rapidly than in width, a scale from the
mid-dorsal region transforming from a structure as wide as long to one
in which the length is four times the width. Old animals with cuspless,
worn, elongate scales present a drastically different appearance from the
animals aptly described by their specific name. (Compare figures 1 and
2, Plate XXXVI.)

Hair changes during life are striking but subject to well-marked
individual variation. At birth and for some time afterward the only

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 657

hair is that of the orbital ring. By the time the animal is half grown the
unscaled parts of the skin are covered with asparse growth of short, light-
colored hair. The usual course of development is that the hair becomes
increasingly dense and long, attaining a length of 20 mm. over the ab-
domen. Some areas, few of which are constant, become a Vandyke
brown. The hair of the arms on adults is always colored in this manner
and any other section may be so. Frequently the sides of the face, the
rear legs, and the perineal region are suffused with this color and the throat
and belly blotched with it.

INDIVIDUAL VARIATION.—Individual variation in length and color
pattern of the hair is treated in the paragraph above. Variation in
color of the scales appears to be little subject to random variation,
though occasional specimens show a few unpigmented scales on the
sides of the body and on the tail, which are probably not post-mortem
changes, since they are seen in anembryo in alcohol. Scale counts of the
transverse longitudinal body rows, of marginal and median caudal seales
have been recorded by Frechkop and earlier writers. Ten specimens
selected at random, with equal distribution among the sexes, show the
following variation in count. My findings are followed in parentheses
by the corresponding figures summarized in Frechkop. It is shown by
my count that variation within one district is great enough to indicate
that scale counts can be of little or no service in recognition of the source
of any specimen of the species.

Number of pre-caudal median scales 27-30

Number of median body scales 19-21 (18-22)
Number of longitudinal rows 21-25 (19-23)
Number of marginal caudal scales 35-40 (34-38)
Number of median caudal scales 29-36 (30-33)

Sex DIFFERENCES.—I am unable to discover external differences
between the male and female tricuspis other than those of the sex organs
and the lacteal apparatus. The males grow to greater length, as may be
seen from the summary of the collector’s measurements. The slight
difference between the ratio of tail to head and body length obtained
from these figures (co 168 per cent—@ 166 per cent) is almost certainly
of no significance.

GEOGRAPHICAL VARIATION.—As suggested above, the external differ-
ences between members of this species from different localities seem to be
inconstant, and whereas I am of the opinion that the animals from
opposite extremes of the range present divergent tendencies, I have not
found a single character which may be relied upon to differ in a constant

658 Bulletin American Museum of Natural History [Vol. LX VI

manner. Thus specimens from the Kasai and from Angola appear to
have longer hair which carries less pigment than that in all but a few
individuals from the northeastern Congo, and this same tendency is
indicated by some of the ten scattered specimens from the Guinea coast
which have come into my hands, though these specimens are too few
and too closely approximated by specimens in the Congo Expedition
series to warrant recognition of races within the species.

CoLLEcToR’s MEASUREMENTS

TotaL LENGTH Tait LENGTH Foot LENGTH
02 Oo, 2 777.9 (617-1027) 463.4 (3850-607) 57.1 (45-75)
25 ot 793.2 (617-1027) 469.6 (360-607) 58.0 (45-75)
Ws © 768.4 (630— 920) 460.4 (850-590) 56.4 (45-75)

CRANIAL CHARACTERS.—Individual variation in the skulls of tri-
cuspts 1s localized and does not approach the extreme variation in general
form found, by contrast, in our much smaller series of longicaudatus.
The features of tricuspis subject to most pronounced variation are the
nasals, the lacrimals, and the foramina. The nasals vary most strikingly
in the form of their lateral wings. These are occasionally bilaterally
asymmetrical, as, for example, in one specimen in which the suture sep-
arating the right and left nasals is neither straight nor in the median
plane. The lacrimals are in some specimens totally within the orbit,
while in others they extend far forward between the frontal and maxillary.
They are in some skulls high above the zygomatic processes of the maxil-
lary and in others extend outward and downward to take some part in
its formation. The foramen magnum and all of the smaller foramina
present a wide range of variation in size and form.

Age variation presents no peculiar features in the species. Unlike
the condition in gigantea, but in agreement with that in longicaudatus,
the sutures of the roof and sides of the cranium seem never to fuse to the
extent of obliteration of the sutures, though they are tight throughout
the period of adult life.

The four occipitals unite early to form a single bone.

The suture of the basal components of the cranium unite in this
sequence from the rear forward: basioccipital-basisphenoidal; basi-
sphenoidal-presphenoidal. Other sutures seem never to close. Even in
the oldest skulls the sutures of the palatal region are widely open.

Sexual differentiation appears not to have occurred in the skulls
except with reference to size. As may be seen from an examination of the
table of cranial measurements, the males exceed the females in size, as
they do in the other African pangolins.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 659

The Ivory Coast representatives may be somewhat distinct from
that population represented by the Congo collection, for there is no
parallel in this latter series for two features seen in a single skull of an
immature specimen from the first locality. The two matters in which this
skull differs are those of size and nasal shape. The posterior border of
the combined nasals is here cuspidate, in contrast to the acute nasals of
the compared series. The skull is also smaller and of denser bone than
skulls of a similar stage of development in the Congo series, and it may
be that in the western limits of the range these features are constant.
This is rendered the more probable by Sjéstedt’s (1897, p. 45) observa-
tion that trzcwspis from the east 1s somewhat larger.

DISTINCTIONS BETWEEN THE SKULLS OF
M. tricuspis AND M. longicaudatus
The classification of the manids has been based chiefly on the ex-
ternal characters, sternae, and vertebrae. Nevertheless skull characters
are well marked, and it may prove useful to note the diagnostic differ-

ences between the two arboreal forms here involved.

tricuspis

1.—The lateral borders of the skull
converge anteriorly in a regular course.
The muzzle is broad.

2.—The mastoid region is lightly in-
flated.

3.—The zygomatic process of the
temporal is large and diverges strongly
outward.

4.—The _ sphenopalatine foramen
opens ventrally and is always prominent
as seen from below.

5.—Slight variation in general skull
shape.

longicaudatus

1.—There is a well-marked anteorbital
constriction, and the muzzle in conse-
quence is narrow.

2.—The mastoid region is strikingly in-
flated.

3.—The zygomatic process of the
temporal is small and only lightly
divergent.

4.—The sphenopalatine foramen
opens posteriorly and is either incon-
spicuous or masked by the palatal flare.

5.—High degree of variation in skull
shape.

Cranial Measurements of Adult Manis tricuspis

Greatest length
Basal length
Palatal length

Breadth across zygomatic

processes

Greatest breadth of brain case
Greatest length of nasals
Breadth of single nasal
Greatest length of mandible

22 F

72.8 (63.8-80.8)
66.8 (57.9-75.4)
41.4 (34.5-44.6)

27.3 (22.7-32.0)
32.3 (28.5-36.3)
27.1 (21.5-32.5)

538 ( S&S 70)
48.4 (41.7-57.0)

Average (minimum-maximum)

20 9

68.7 (58.5-79.2)
63.1 (51.6-73.3)
37.0 (31.5-43.3)

25.4 (20.2-29.3)
30.6 (27.7-32.9)
24.3 (18.1-31.2)

4.4 ( 3.5- 7.0)
45.7 (37.8-53.1)

44 3, 9
70.7 (58.5-80.8)
64.9 (51.6-75.4)
39.0 (31.5-44.6)

26.4 (20.2-32.0)
31.5 (27.7-36.3)
25.7 (18.1-32.5)

5.0 ( 3.5- 7.0)
46.3 (37.8-57.0)

660 Bulletin American Museum of Natural History [Vol. LX VI

ReEpRoDucTION.—Two embryos from the Congo collection are at
hand. One of these was taken from a specimen 810 mm., in total length,
killed at Medje, January 24, 1910. This embryo was 80 mm. long. A
female taken November 16, 1909, carried an embryo 280 mm. long,
which is but 22 mm., shorter than another specimen secured alive on
December 4, at Lukolela by another expedition. It is indicated, then, that
the young are approximately 290 mm. long at birth, and that they are
sometimes born in November or December.

There is usually a single pair of mammae, as noted by the ‘allen
and as may be seen from examination of dried skins. A single case was
recorded in the field catalogue as ‘“‘two pairs pectoral mammae,’”’ but I
am unable to discern more than a single pair on the skin of this same
specimen.

Movements.—The field notes on the feeding habits of this species
were included in the Bequaert report. Mr. Lang’s other notes concern-
ing locomotion were made from observations on four individuals. These
notes are transcribed from the field catalogue with slight alterations and
paragraphed separately.

No. 1.—This specimen was an expert climber and surprisingly rapid in her
movements. On the ground she would cover at least four feet in a second, shuffling
along in a peculiar manner. When taken by the tip of her tail she would at once
hook the claws of her hind limbs into the large lateral scales near the base of her tail,
and either ascend by climbing up or, by bringing her body to a position at a right
angle to her tail, she would swing around hooking after everything her fore limbs could
possibly reach. In this position her horizontal body could make a three-quarter twist.
She could then look toward the ground and halfway up again toward the sky. She
would roll up when touched on her snout and would do this so rapidly that one could
hardly take one’s hand away quickly enough to avoid getting caught between the
scales.

No. 2.—When ascending or descending a tree of about four inches in diameter
this manis formed a half circle with the end of his tail, the lower surface of which was
kept in close touch with the bark. In going up and down oil palms he moved his tail
sidewise and up and down, probing for and taking hold of the slightest prominence
with the tip of his tail. The skinny pad on the lower side was evidently of great
sensibility. Once he had taken hold of any prominence or small branch, he at once
worked freely with his fore limbs, as he could hold and direct himself when fastened
only by his tail. As a rule, however, he used his hind limbs in keeping any position
lasting more than a moment.

No. 3.—This young manis was still alive. Put upon the back of a dead though
still unskinned male, it at once climbed toward the tail, grasping it firmly on the
edges, and put its tail around the tail of the adult specimen. When the adult speci-
men was shaken the young seemed not to be troubled in the least. The trial was re-
peated several times, and every time it selected the tail to which to attach itself, some-
times with the head forward, sometimes backward. It is quite certain that the young

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 661

travel clasped firmly on the upper side and parallel with the tail (see Plate XX XVII,
figure 1). It would roll itself like the adult specimens.

No. 4.—It was able to shuffle along as rapidly as a man can walk, always touch-
ing the tips of its claws to the ground, on the forelegs as well as hind legs. The manis
rolled up whenever touched, hooking even the tip of its tail over the dorsal scales.
When shaken by the tip it unrolled at once and started to walk. Sometimes the
animal sat on its hind legs, and the forelegs then nearly touched the ground. It
climbed well on trees and also on steep ground. It could hold itself as firmly with its
tail as with its forelegs or hind legs alone. The animal never tried to scratch with its
claws. Its only defence consisted in rolling up, though in a very short time it un-
rolled again (Plate XXXVI, figure 2) and walked away. Though evidently noc-
turnal, it seemed not to mind the sun at all.

ORYCTEROPODIDAE

OrycrTERoPus Geoffroy

Orycteropus GEOFFROY, 1795, ‘Décade Philosophique 1795.’ [From Agassiz

1842, ‘Nomenclature Zool.,’ Mamm., p. 23.]
Type.—M yrmecophaga capensis GMELIN.

Of the thirteen aard-varks secured by the Congo Expedition two, an
old male and an old female, came from Niapu, well within the forest.
Eleven others, ranging from a very young animal with unworn teeth to
mature individuals of both sexes, came from the savanna region of
Faradje. Because of the lack of sufficient comparable material and the
insignificant diagnoses given most of the sixteen nominal races, it has
proved difficult to ascertain the affinities of the animals in the collection.
The two old animals from the forest present many characters by which
they may be separated from those of the savanna, 350 kilometers (i.e.,
c.217 miles) to the northeast, but some of these appear to be age charac-
ters, and there are relatively few that can not be attributed to senility
or random variation. These few, however, are well marked and have
appeared to be of sufficient importance to warrant the recognition of
two races. L6nnberg’s description of O. erikssoni from Guffuri (Gufuru)
on the lower Bomu River agrees closely with the Niapu (forest) race,
whereas the points in which the Faradje specimens approximate the
characterization of erzkssoni are, with two exceptions, those common to
both Niapu and Faradje specimens. The two exceptions mentioned are
the size of the praemaxillae (as measured by dividing the distance from
the maxillo-praemaxilla-nasal juncture to the anterior end of the prae-
maxillae into the nasal suture) and the size of the teeth. In the first
exception the praemaxillary measurement is half or less than half the
length of the nasal suture, whereas this measurement in the forest speci-
mens is well over the nasal length. In the eleven specimens from the

662 Bulletin American Museum of Natural History [Vol. LXVI

savanna country I find no evidence that the praemaxillae increase dis-
proportionately in late life, and I conclude that in this aspect the Niapu
specimens are not perfectly typical of erzkssonz. The one other likeness
of the savanna specimens to the type of erzkssonz lies in the diameter of
the teeth which are noticeably larger than those of the forest specimens.
Thus longitudinal and transverse diameters of the next to the last molar
in the largest animal from the forest are 10.0 and 6.0 mm., respectively,
whereas corresponding measurements in a large savanna specimen are
12.5 and 9.0, exactly the measurements reported for erzkssoni. This
skull is, however, much smaller than that of erzkssont (24 mm.
shorter). The series at hand offers faint suggestion that in senility
the diameter of the teeth decreases with the total number. If this is
true the small teeth of the two Niapu aard-varks (and Hirst’s Cameron
race, leptodon) are not diagnostic of race but only of age.

It is concluded then that the forest race of the northeastern Congo
may be referred to Lénnberg’s erzkssonz. From available descriptions I
was not able to refer the aard-varks from Faradje to any of the described
forms, though on geographical grounds it would seem possible that they
might resemble kordofanicus Rothschild. The meager note accompany-
ing the proposed name for that race states that, with certain exceptions
(which may be juvenile characters), the type is nearest aethiopicus,
from which the Faradje aard-varks differ in several respects. I have
therefore considered it necessary to propose a new name for the race from
Faradje, which has been given as Orycteropus erikssoni faradjius Hatt,
in a preliminary report.!

Orycteropus erikssoni erikssoni LOnnberg
Figure 2a

Orycteropus erikssont LONNBERG, 1906, Arkiv for Zoologi, Stockholm, Bd. III,
INOS Sy JDs Ike

Orycteropus e. ertkssoni is a large aard-vark occurring in the forests of
the Congo. It is characterized by short hair, short ears, and large claws.
The skull is large, the anterior base of the zygoma narrow, and the
mandible slender.

Represented by an old & and an old @, skins with skeletons, col-
lected at Niapu, November 24 and December 3, 1913.

The collector’s measurements of these animals are:

11932, Amer. Mus. Novitates, No. 535, p. 1.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 663

The two skins show the same sex dichromatism described below for
O. e. faradjius but do not exhibit any coat character by which I can
distinguish them from this other race.

The ears are very short and measure from notch to tip on the tanned
skins 95 and 93 mm., in contrast to corresponding measurements of 120,
120, 125, and 140 mm. on the four largest specimens from Faradje.

The claws of O. erzkssoni are larger than those of O. e. faradjius,
as shown by measurements of the claw of the third digit of the manus.
In the former, this length, from eponychium to nail tip is 54 mm. in one
of the two specimens, 55 in the other. Similar measurements on four
adult O. e. faradjius are 40, 40, 43, and 45 mm.

The skull of Lonnberg’s aard-vark is easily distinguished from that
of Faradje specimens by size, the character of the anterior base of the
zygoma, the mandibular proportions, and the less conspicuous features
noted in the diagnosis of the savanna race. The species differs unmis-
takably from such Abyssinian, East African, and South African animals,
as I have seen in many features, of which, as Grote has shown, the man-
dibular are among the most striking.

Orycteropus erikssoni faradjius Hatt
Plates XX XVII, Figure 2; XXXVIII; XXXIX; Figure 26
Orycteropus ertkssoni foradjius Hart, 1932, American Museum Novitates No.
5395p: L.

Orycteropus e. faradjius is a large aard-vark occurring in the savannas
of the northeastern Congo. It differs from O. e. erzkssonz in smaller size,
longer ears, and shorter claws. The anterior base of the zygoma is
broad (over 20 mm.), the mandible heavy.

Orycteropus e. faradjius is represented in the collections of The
American Museum of Natural History by 9 skins, 11 skulls with 10
skeletons, collected at Faradje between May 27, 1911 and February 24,
1913. Of these eleven individuals 4 are adult males, 6 adult females, and
one a juvenile male.

The collector's measurements of the adults as summarized into
averages, minima and maxima, are:

ToraL LENGTH Tart LenetH Foor LenetH Ear LENGTH

Oot, 1750 (1610-1850) 642 (580-720) 273 (263-290) 170 (158-180)

4c 1721 (1640-1785) 637 (610-720) 276 (270-290) 169 (162-175)

Gao 1770 (1610-1850) 644 (580-700) 270 (263-275) 171 (158-180)
The height at shoulder of two adult females was recorded as 400
and 370 mm., respectively. Other measurements of a female whose total

Fig. 2b
Fig. 2. a.—Skull of Orycteropus e. erikssoni, adult &, Niapu, Belgian Congo (No. 51875); 6.—Skull
of Orycteropus e. faradjius, adult o&, Faradje, Belgian Congo (No. 51373). Type specimen.

664

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 665

length was 1780 were: length of mouth from tip of lower Jaw to corner
of mouth, 52; from middle of eye to end of snout, 185.

This aard-vark differs externally from O. e. ertkssoni in its slightly
smaller size, longer ears, and shorter front claws. The skull differs from
erikssoni, as represented by the two specimens from Niapu and the type
description, in the following features: The inflation of the region of the
frontonasal suture is more pronounced, and, in consequence, the upward
curve of the posterior part of the nasals is greater. The nasal region is
also broader, to the extent that the greatest width of the combined nasalia
is more than one half the length of the nasal suture. The lacrimal bone
of faradjius is comparatively long, and the length of its suture shared
with the frontal anterior to the orbit is about 70 per cent of the total
length of the lower frontal border between the orbit and the posterior
external angle of the nasal. In erzkssoni the lacrimal-frontal suture is
60 per cent or less of the length of the lower frontal border. The zygo-
matic arch at the level of the end of the zygomatic process of the maxil-
lary is very broad (over 20 mm.), while in erzkssonz, the arch is very
narrow, its depth measuring less than 20 mm.

The mandible of faradjiuws is more massive and broader, and the
angular and coronoid processes are higher than in erzkssonz. Thus the
greatest height of the mandible equals about 45 per cent of its greatest
length in the race from Faradje, whereas in erikssonz this height is only
39 per cent of the length. The greatest breadth of the mandible in the
former race is contained in the mandibular length 4.8 times, whereas in
the latter race the length is near 5.5 times the breadth. The tip of the
angular process lies midway between the alveolar plane and the articular
process, whereas in the larger forest form it is much lower, the height
above the alveolar level being only about one half the distance from the
tip of the angular process to the tip of the articular.

SEXUAL DIFFERENCES.—Sexual dichromatism, which seems not to
have been noted in the literature, is well marked in adult specimens. In
the males the hair of the entire head is dark brown, but in the females
that below the orbits, except for a dark gular stripe, is light-colored. On
the outer surface of the thigh the hairs are slightly blacker in males than
in females, in which sex the hairs retain more of the reddish wash of
youth.

Age change of color in both sexes involves an intensification and
extension of the pigmented areas of the head (particularly in the males),
a progressive blackening of the dark area of the limbs and shoulders
that all but obliterates the deep red color found on immature specimens,

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668 Bulletin American Museum of Natural History [Vol. LXVI

and, in contrast to these changes, a loss of virtually all pigment in the
hair of the back, so that this area changes from one of deep bay, mod-
erately long hairs to one of short, dirty-white hairs. The hairs of the
distal two-thirds of the tail are uncolored throughout life, though the
underlying dermal pigment, particularly of the basal half of the tail,
may in the tanned skin, give a mottled pattern of yellow and gray.

Sex differences in the skull are not found in this series, though from a
comparison with other material it seems probable that males attain
greater size than do the females.

AGE DIFFERENCES.—Age changes in the skull are those of size, suture
obliteration, and proportion. ‘The first sutures to be obliterated are
those separating the occipital elements, the parietal suture, and, later,
the suture between the interparietal and parietals.

This later suture is open in a skull 250 mm. in greatest length,
whereas the others, which were open in one of 181 mm., are completely
obliterated in this larger skull. In skulls of yet older animals the parietal-
interparietal suture is closed. The occipitoparietal suture, however, is
open in the oldest specimens. Later the frontal suture closes progres-
sively from the rear. Fusion occurs along the nasal suture very sporadi-
cally, being found only in two young skulls. The most prominent modi-
fications of skull form are a great reduction in the proportionate degree
of inflation in the anterior frontal region, and marked increase in skull
length in proportion to skull depth.

The teeth commonly regarded as the first and second molars, when
first erupted have two cones, but these are soon lost through wear. The
skull of a newborn individual on which these cones are displayed, also
shows double cones on the last premolar. On the left side two cones are
also present on the next to last premolar, while on the right the area is
occupied by two simple teeth which suggest a splitting of germinal
buds as the explanation of the large number of maxillary teeth reported
for the genus. The diameter of the teeth increases until maturity, but
with approaching senility the anterior teeth are lost, and the posterior
members may be slightly reduced in size.

Races then, the characters of which are founded on color, size,
degree of frontal inflation, and tooth diameter must be critically ex-
amined with reference to the age class of the type.

The skull measurements of the adult specimens are represented in
their averages, minima, and maxima in preceding table, pp. 666-667.

Firtp Datra.—Mr. Lang’s field notes on specimens of this race
contain observations on anatomy, habits, and habitat, which may be
conveniently arranged according to subject matter.

1934] Hatt, American Museum Congo Expedition Pangolins and Aard-Varks 669

EXTERNAL CHARACTERS.— The ears of an adult female were whitish
pink on the inside. The irises of this and another individual were a very
dark brown.

The skin, particularly on the neck, was arranged in tiny folds.
There was a pouch on either side of the vulva that contained a strongly
smelling matter.

There are two pairs of abdominal mammae.

Digestive Tract.—In an individual 1780 mm. in total length the
small intestine measured 4520 mm., the large intestine 2450, and the
caecum 130 mm. The digestive tract at the base of the latter was
greatly enlarged. Near the pylorus was a large muscular gland.

Foop.—The stomachs of five contained termites, so well chewed up
that none were complete.

LocoMOTION AND FrEepING.—Mr. Lang wrote that the aard-vark
usually walks on tiptoes, though it may progress for a few steps with the
whole hind foot resting on the ground. When walking, the nose is kept
close to the ground, and then the nostrils are opened and shut by con-
tracting the skin muscles pulling from the rear. The tail tip touches the
ground when the animal is moving. The animal frequently sits on its
haunches and when digging assumes this position. The tail, then, bears
considerable weight.

The thick heavy claws borne by the muscular fore limbs are nearly
straight, and with these chisel-like points the aard-varks have no difficul-
ty in breaking into the hard, solid structures of the termites. These
animals are licked up by the hundreds with the long slimy tongue as
they assemble quickly in large masses near the point of destruction,
either to defend their home or to repair it.

Burrows.—The field book carries the following note on burrows:
“The Orycteropus live in burrows (probably in small families or singly)
which they dig themselves in dry, hard ground. The excavated ground
is thrown up in small hillocks close to the opening. These burrows are
fairly large, the size of the tunnels allowing a small man to enter without
great difficulties. Of seven burrows I have seen, two had only one
entrance, four had two and one had three. The entrances of one tunnel
were about 14 meters from one another at the maximum. All these
burrows were deserted, and it is probable that they inhabit several
burrows alternately, and it is sure that they occasionally dig a hole
simply for shelter.” (Plate XX XVII, figure 2.)

Time or Activiry.—‘ The Orycteropus,” observes Mr. Lang, “‘is
rather difficult to procure, not only an account of its nocturnal habits,

670 Bulletin American Museum of Natural History [Vol. LXVI

but principally by reason of its scarcity. They roam about during the
night, singly or in pairs, as may be seen from the impressions of their
claws left on moist ground.”

Hasitat.—It was recorded that an aard-vark was captured by
natives somewhat north of Rungu. North and northeastward of
Niangara to Faradje the animal was said to occur everywhere. Mr. Lang
considered it a ‘“‘constant resident of the vast plainlike stretches which
are abundantly covered by stunted trees and thick low bushes.”’ There
termites were abundant, and as they constituted the only food of Orycter-
opus, these animals found an ample supply and were usually well
nourished.

HuntTING BY THE Natives.—The relation of the aard-vark to the
native life in the district as it existed at the time of the Congo Expedi-
ton is suggested by the following extract from the field book: ‘The
native manner of securing the Orycteropus in these regions lends itself
to adventure and has given rise to general and very deep set superstitions.

“Tt is currently held that some have lost their lives in the burrows
and have never been heard of again. The native hunters usually track
the Orycteropus to its burrow after a heavy rain, and having ascertained
the exact locality they start out in numbers to secure the highly valued
prize. Among the Azande and Logo, and probably among related
tribes, it is the custom that one young fellow well supplied with ‘medicine’
enters the burrow armed only with a short handled spear or a long knife.
If he is lucky he encounters the beast which usually tries to save itself by
digging and throwing the excavated ground in the face of its pursuer;
if it succeeds in placing a wall between the man and itself before he can
successfully place his spear the man considers the undertaking a failure
and tries to retreat. If however, the native can kill it he will indicate
his position to the eagerly waiting outsiders by tapping against the
upper wall. As rapidly as possible the natives sink a shaft in his direc-
tion and lift out the dead Orycteropus and the valiant hunter who is
seldom deeper than five feet below the surface.

“The teeth are worn around the wrist by the Mangbetu, Azande,
Logo, and affiliated tribes to prevent illness and to ward off ill fortune.
The bristly hair of the nostrils and that between the toes is said to be
fatal if well powdered and thrown into their beer. The consumer’s neck
is said to swell to such an extent that death usually occurs after three
days. The claws carried in the baskets of women that are collecting the
winged edible variety of termites insure an ample supply. The meat has
a strong odor and the appearance of pork; it is eaten by the Azande,
Mangbetu, and Logo.”

BIBLIOGRAPHY

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des Pangolins et de leur utilisation en Taxonomie.’ Bull. Mus. Nat.
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1919b. ‘A propos de la Taxonomie des Pangolins: Rectification au Régne
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CHRISTY, CUTHBERT. 1924. ‘Big game and pygmies.’ 325 pages, London.
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FRECHKOP, SERGE. 1931. ‘Quelques observations sur la classification des Pangolins
(Manidae).’ Bull. Mus. Roy. d’Hist. Nat. Belg., Tome VII, No.
22, pp. 1-14.

Gray, J. E. 1865. ‘Revision of the genera and species of entomophagous Edentata,
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seum.’ Proc. Zool. Soc. London, pp. 359-386.

1873. ‘Handlist of the edentate, thick-skinned and ruminant mammals.’

176 pages, London.

GrRoTE, HERMANN. 1921. ‘Neue Erdferkel (Orycteropus) aus Deutsch-Ostafrika
und Kamerun.’ Archiv for Naturgesch., Band 87, Abt. A, Heft
7, pp. 121-127.

Hirst, A.S. 1906. ‘A new species of Orycteropus.’ Ann. Mag. Nat. Hist., (7) XVII,
pp. 383-384.

JENTINK, F. A. 1887. ‘Catalogue Ostéologique des Mammiféres.’ Mus. Hist. Nat.
Pays-Bas, IX, 360 pp., 12 pls.

1882. ‘Revision of the Manidae in the Leyden Museum.’ Notes from the
Leyden Museum, IV, pp. 193-209.

LONNBERG, E. 1906. ‘On a new Orycteropus from northern Congo, and remarks on
the dentition of the Tubulidentata.’ Arkiv fér Zoologie, Stock-
holm, ITI, Art. 3, pp. 1-35.

671

672 Bulletin American Museum of Natural History [Vol. LXVI

Martscuig, P. 1894. ‘Die naturliche Verwandtschaft und die Verbreitung der
Manis-Arten.’ Sitzber. Ges. natur. Freunde, Berlin, No. 1, pp. 1-13.

Pocock, R. I. 1924. ‘The external characters of the pangolins (Manidae).’ Proc.
Zool. Soc. London, pp. 707-7238.

RAFINESQUBE, C. 8. 1821. ‘Sur le genre Manis et description d’une nouvelle espéce:
Manis Ceonyx.’ Ann. Génér. Sci. Phys. Brux., VIT, pp. 214-215.

ROTHSCHILD, WALTER. 1907. ‘Ona new race of Orycteropus.’ Nov. Zool., Tring,
XIV, p. 506.

ScHOUTEDEN, H. 1930. ‘Les Pangolins.’ Rev. Zool. et Bot. Africaine, XVII (4):
Bull. Cercle Zool. Congolais, VI, (3), pp. [87]-[95].

Scuusotz, HERMANN. 1912. ‘Zoologische Beobachtungen wahrend der II Wissen-
schaftlichen Innerafrika-Expedition S. H. des Herzogs Adolf
Friedrich zu Mecklenburg, 1910-1911.’ Ber. Senckenb. Natur-
forsch. Ges. Frankfurt am Main, XLIII, pp. 324-358.

SonntaG, C. F., Woouuarp, H. H., anp Cuark, W. E. LE Gros. 1925-1926. ‘A
monograph of Orycteropus afer.’ Proc. Zool. Soc. London, 1925,
pp. 331-487, 1185-1235; 1926, pp. 445-485.

SUNDEVALL, C. J. 1845. ‘Uebersicht der Gattung Manis.’ Isis, XX XVIII, pp.
583-588.

THILENIUS, GEoRG. 1912. ‘Uebersicht ueber die wissenschaftlichen Ergebnisse der
deutschen Zentralafrika-Expedition 1910-1911.’ Adolf Friedrich
Herzog zu Mecklenburg’s ‘Vom Kongo zum Niger und Nil.’
Leipzig, II, pp. 367-384.

Tuomas, OLDFIELD. 1890. ‘A milk dentition in Orycteropus.’ Proc. Roy. Soc.
London, XLVII, pp. 246-248.

PDNTIS LOCO no LOOKER

Pirate XXXII
Fig. 1. Manis gigantea. Young female, Poko, August 10, 1913 (No. 53860).
Total length, 810 mm.

Fig. 2. Manis gigantea. Adult female, Niangara, April 26, 1913 (No. 53846).
Total length, 1360 mm. Note the uneven wear of the scales, indicative of advanced
age.

oL. LX VI, Puate XXXII

Buuuetin A. M. N. H.

Puate XX XIII
Fig. 1. Manis gigantea. Adult female, Niangara, April 26, 1913 (No. 53846).

Fig. 2. Manis gigantea. Young female, Poko, August 10, 19138 (No. 53860).

IWIXXX 41d ‘TAX'[ “LOA “HON IW ‘V Niwaring

PuaTE XXXIV
Fig. 1. Manis gigantea. Embryo. Niapu, December 9, 1913 (No. 53856).
Note the keels on the scales of the lower flank and the rounded contours of the marginal caudal
scales. The nipple, visible on the flank, is about one half the size of that near the axilla, which, in this

photograph, is hidden by the arm. The claws in this fetus are bifid, as are the ungual phalanges of the
adults.

Fig. 2. Burrow of Manis gigantea, near Babeyru, July, 1914.

Vou. LX VI, Puate XXXIV

Buuuretin A. M.N. H.

Fig

PLateE XXXV
Fig. 1. Manis longicaudatus. Adult male, Niapu, December 16, 1913 (No.
53871).

Fig. 2. Manis longicaudatus. Adult female, after death, Akenge, October 10,
1913 (No. 53862).

AXXX 3LV1q ‘IAXT “TOA ; ‘HN CIN CV Nivaiiag

Pirate XXXVI
Fig. 1. Manis tricuspis. Juvenile female, Avakubi, October 29, 1909 (No.
53888). This individual is a good example of the age period in which the dorsal scales
are tricuspid.
Fig. 2. Manis tricuspis. Adult male, Stanleyville, August 9, 1909 (No. 53938).

The mid-dorsal scales of this individual are no longer tricuspid, though this pattern is still retained
on tne scales of the head and flank. When photographed the animal was not so closely coiled as is
possible.

IAXXX @14V1g ‘TAXT “TOA "HN IN ‘VENI TING

PuaTts XXXVII

Fig. 1. Manis tricuspis. The young of this species are transported in the
manner here shown (see p. 660).

Fig. 2. Burrow of Orycteropus ertkssoni faradjius. Faradje, March 6, 1911.

ot. LXVI, Prats XXXVIL

Butuetin A. M. N. 4H.

ig.

See

PuaTeE XXXVIII

Fig. 1. Orycteropus ertkssoni faradjius. Adult female, Faradje, March 6, 1911
(No. 51235).

Fig. 2. Right manus of Orycteropus ertkssoni faradjius.

Fig. 3. Left pes of Orycteropus ertkssoni faradjius.

Vou. LX VI, Puate XX XVIII

N. H.

Butuetin A. M

Puate XXXIX

Figs. 1 and 2. Orycteropus ertkssoni faradjius. Faradje, March 6, 1911 (No.
51235).

The long ears, short claws, and inflation of the frontonasal region characteristic of this subspecies
are well illustrated by this specimen.

XXXIX

=
D}

Vou. LXVI, Puati

Butuetin A. M. N. H.

Fig. 2

Fig. 1

5 an

We)
Leer A a
i

Hyraxes Collected by the American Museum » Congo
Heres

By ha T. Harr

BULLETIN
OFr

THE 5 AMERICAN MUSEUM OF NATURAL eo ae.
‘You. LXXII, Art. IV, pp. 117-141
New York

Issued August 28, 1936

Article IV.—THE HYRAXES COLLECTED BY THE AMERICAN
MUSEUM CONGO EXPEDITION!

By Rosert T. Hatr

Puates XVIII ro XXII; Text Figures 1 To 3

PAGE
1 AT TTTE OTB GETING, Aig EoRESt oo cle o lec geneeeeo e tls oet Ae Lal?
SPECIES WITH THEIR LOCALITIES AND NUMBER OF SPECIMENS FROM EACH

TEXOVGUNTLTETENS, os Sow ed Se neg Ae ee cy es ICR ter Oe Ree ee ea ee 119
List or LOCALITIES, WITH NAMES OF THE SPECIES AND SUBSPECIES AND NuM-

BUR LO SPECGIMENG (bAKEN AT WACH LOCALIMY 45 5225.45.06. 5525505 oo 119
MARION NOE INDICATIVE OF RELATIONSHIP... 26.0... 5 oh eds es on ew oe ns 119
Some CHARACTERS UTILIZED FOR CLASSIFICATION OF HYRACOIDEA AND THEIR

WINACRION: dN REE 1 CONGOUSPECIES=. jis 5 0) fo 5 oie eh sak a ienenn 124
OW GOL: CORSGIUS ONT OLOT «oa 8 aie fe. p22 ee Ws I og Reds oo hale God OS) Sone 126
TARO RELL AOLSOUISNCMIND Sey pete Rahen Alem eA ole Ae wee) ole Mare 2 » Ble
ENG DILUTE CDCI Os ea pa RRS Se adage Tra ee care a elt32
NaC CRD IOR OIMUSIONMMOPESU ANA. taints sae yaa ace eee eee te 3a son 135

INTRODUCTION

The American Museum Congo Expedition, of which Mr. Herbert
Lang was leader and Mr. James P. Chapin assistant, operated in the
Belgian Congo from 1909-1915.2 Among the excellent series of mam-
mals obtained were ninety-eight hyraxes. Particularly well repre-
sented was the then little-known Emin’s tree hyrax. A novelty was a
new species, Heterohyrax chapini, described in a preliminary report.®

Study of the Hyracoidea of the collection was initiated in 1932,
but the manuscript left uncompleted pending the appearance of Hahn’s
monograph? of this group. The collection on which the present report
is based represents rather fully the hyrax fauna of the northeastern
Congo, and thus furnishes a good basis for the appraisal of Hahn’s
conclusions concerning the taxonomy of the species present in this
same area.

1 Scientific results of the Congo Expedition, Mammalogy, No. 17.

2 A general account of the expedition with a map of its collecting grounds may be found
under the following reference:

Osborn, Henry Fairfield, 1919, ‘The Congo Expedition of The American Museum of Natural
History,’ Bull. Amer. Museum of Natural History, X X XIX, pp. xv—xvii.

3 Hatt, Robert T., 1933, ‘An Annotated Catalogue of the Hyracoidea in the American Museum
etc.,, American Museum Novitates, No. 594, 13 pages.

pion Herbert, 1935, ‘Die Familie der Procaviidae,’ Zeitschrift fiir Siugetierkunde, Bd. IX,

pp. —358,

118 Bulletin American Museum of Natural History [Vol. LX XII

Hahn’s monograph, a long-needed review of recent and fossil Hyra-
coidea, is based on the excellent collections in Berlin, Frankfort and
Tervueren, together with notes compiled by the late Doctor Brauer on
collections in other continental museums. It is unfortunate that the
author had no opportunity to examine the British Museum Collection
nor any of the material in America, specimens which are of im-
mense importance in any study of this group of animals. Included
in the work is a sound discussion of the skull and skeletal characters,
used as criteria of relationship, together with a dissertation on the
ecological and zoogeographical background of hyrax phylogeny.

Most of Hahn’s conclusions appear sound, particularly as regards
the main phyletic trends. As may be seen in the following treatment
there are, however, numerous points on which I cannot agree with him
concerning the taxonomy of Congo forms. ‘Thus, it appears to me highly
probable that Dendrohyrax d. latrator must be recognized; and I am
convinced that there is no basis, other than Hahn’s failure to recognize
dichromatism, for considering the tree hyraxes of the northeastern Congo
as anything but homogeneous. The fact that in the monograph my
Heterohyrax chapini was listed under another species was due obviously
to the fact that my previous report on the Order did not appear until it
was too late for Doctor Hahn to give it consideration. The omission in
Hahn’s revision of all reference to types and type localities is an unfor-
tunate neglect, particularly in that his predecessor, Brauer, so fre-
quently made no mention of the number, sex or age of his types. Ap-
parently overlooked in the review was Dendrohyrax rubriventer Brauer
(= D. d. eminit Thomas).

It is again my pleasure to tender my thanks to Doctor James P.
Chapin for his invaluable advice; and to the authorities of the Field
Museum, the United States National Museum, the Museum of Compara-
tive Zoélogy and the British Museum for their courtesy in permitting the
examination of collections in their charge. |

The text figures in this Bulletin were drawn by Marcelle Roigneau
Hatt and the photographs of animals in the flesh made by Mr. Lang.

1936 | Hatt, Hyrazxes Collected by American Museum Congo Expedition 119

SPECIES WITH THEIR LOCALITIES AND NUMBER OF SPECIMENS
FROM EACH LOCALITY

SPECIES LOCALITIES SPECIMENS
1.—Dendrohyrax dorsalis latrator _ Bolobo, 1. 1
2.—Dendrohyrax dorsalis emint Akenge, 3; Avakubi, 1; Gaman-

gui, 4; Medje, 6; Ngayu, 2;

Niangara, 3; Niapu, 24. 69
3.—Heterohyrax chapini Matadi, 2. 2
4.—Procavia johnstoni lopest Aba, 24; Faradje, 1; Vankerck-

hovenville, 1. 26

LIST OF LOCALITIES, WITH NAMES OF THE SPECIES AND SUBSPECIES
AND NUMBER OF SPECIMENS TAKEN AT EACH LOCALITY

LOCALITIES SPECIES AND SUBSPECIES NUMBER OF SPECIMENS
Aba Procavia 7. lopest 24
Akenge Dendrohyrax d. emint 3
Avakubi Dendrohyrax d. emini 1
Bolobo Dendrohyraz d. latrator i
Faradje Procavia 7. lopest 1
Gamangui Dendrohyrax d. emini 4
Matadi Heterohyrax chapiniz 2
Medje Dendrohyrax d. emint 6
Ngayu Dendrohyrax d. emini 2
Niangara Dendrohyrax d. emini 3
Niapu Dendrohyrax d. emini 24
Vankerckhovenville Procavia 7. lopest 1

VARIATION NOT INDICATIVE OF RELATIONSHIP

INDIVIDUALITY IN SKULLS

Skulls of the hyraxes show a range in variation in general skull type
beyond that encountered in many mammals. Comparing specimens of
equal apparent age,! as judged by suture closure, development of skull
ridges and dental wear, there are found skulls that contrast to the ex-
tremes shown by the measurements given below for Dendrohyrax d.
emint. Specimens of every intermediate size and shape occur in the

1 For the convenience of the reader, Thomas’s (1892, P. Z. S., p. 53) classification of the de-

velopmental stages of the hyraxes is repeated.
tage I. Before the milk dentition i is fully in place.

Stage IT. Milk dentition all up and in use. M1! not visible.

Stage III. M!tup; M2? below level of bone.

Stage IV. M2? just appearing or partly up

Stage V. M? nearly or quite up, M3 below level of bone.

Stage VI. Tip of M3 appearing.

Stage VII. M2 partly or nearly up, but still unworn.

Stage VIII. M2 up and in use.

120 Bulletin American Museum of Natural History [Vol. LX XII

series and one cannot admit the cases at one extreme or the other as
dwarfism or gigantism.

Cranial Individuality in Adult Males of Dendrohyrax d. emini

Old VIII Early VIII
American Museum Number 53836 53837 53823 53822
Greatest length 2288 110.0 114.5 106.5
Condylobasal length 120.7 110.0 114.0 106.5
Zygomatic breadth 67.5 65.2 63.2 62.3
Anterior edge of orbit to gnathion 43.7 38.5 40.3 34.0
Frontal suture, length oR eAw2 32.9 On 1/ 35.0
Bregma to occiput 49.5 40.0 40.0 40.0
Palatal length 63.5 58.2 60.0 59.0
Basisphenoid plus basioccipital, length 41.2 34.3 35.9 32.1
Premolars plus molars, length 42.3 39.5 Bil ol 41.0

In the instance of the older skulls the two specimens contrasted
differ chiefly in the measurements of length, the discrepancies between
the two being greatest in the postorbital region, but also pronounced in
the preorbital. The two younger skulls, on the other hand, show a
striking difference in the length of muzzle but little difference other than
this.

Similar broad divergence is observed in the Congo Expedition series
of Procavia j. lopest where the skulls of three female specimens ‘all in
Stage VIII and with nearly the same degree of tooth wear (see ‘“‘height
of crown’’) are of three types, one (No. 53776) long and slender, another
(No. 53777) long and broad, a third (No. 53796) short and slender.
The degree of difference is perhaps best shown by the following table of
measurements.

Cranial Divergence of Stage VIII Females of Procavia j. lopesi

American Museum Number 53776 53777 53796
Greatest length 103.0 98.8 92.0
Condylobasal length 102.5 95.7 89.8
Zygomatic breadth 57.2 57.8 51.8
Width at postorbital processes 35.6 40.5 34.0
Length of nasal suture 200 24.4 24.5
Frontal suture, length 39.5 38.4 34.0
Bregma to occiput 33.2 32.7 SH UST h
Diastema, length 15.0 11.6 10.7
Premolars plus molars, length 42.3 41.8 39.3
Height of crown, M3 4.5 4.1 4.1

1936 | Hatt, Hyrazxes Collected by American Museum Congo Expedition IPA

FONTANEL AND WoORMIAN BONES

Fontanel and wormian bones occur with relative frequency in the
Congo Expedition series of Procavia 7. lopesi. None were found in the
skulls of Dendrohyrax d. emini, the early closure of the cranial sutures
being unfavorable to their recognition, even should they be present in
early life. In this respect these two species are representative of their
generic groups, for fontanel and wormian bones are about four times as
frequent in occurrence in Procavia as in Dendrohyraz.

The hypothesis advanced by Schultz (1923, Journal of Mammalogy,
IV, p. 65) that these bones are neomorphs in the mammalia that appear
at places in which the normal roofing-over process is inadequate to meet
demands of protection, receives some slight support from these findings,
for in the hyraxes these accessory ossicles occur with greatest frequency
in the group with the most retarded solidification of the skull roof, and
most rarely, if one may judge from examination of a series that contained
few skulls with open parietal sutures, in the group in which early
closure is the rule.

The incidence of occurrence in the three genera, as determined from a

survey of all skulls of hyraxes in the American Museum, is as follows:
NUMBER OF
SKULLS WITH
WoORMIAN OR

ToraL NUMBER FONTANEL
GENUS OF SKULLS BONES INCIDENCE
Procavia 53 8 1H UY,
Heterohyrax 34 3 8.8%
Dendrohydrax 78 3 B.0%
All forms 165 14 7.8%

The two species of Congo hyraxes represented by series show slightly
different percentages than do the larger groups considered. Thus Pro-
cavia 7. lopesi has five cases of accessory roofing bones in 22 skulls,
an incidence of 30 per cent, whereas no case (unless a single Stage I skull
in which the interparietal bears a median suture should be considered an
exception) occurs in 37 skulls of Dendrohyrax d. emini.

The sexual distribution of extra bones in the mid-line was approxi-
mately equal, there being 6 males and 7 females showing the variant.

ASYMMETRY

Asymmetry is not a character of the Hyracoidea and no case of well-
marked asymmetry in the skulls of Congo hyraxes has been encountered

122 Bulletin American Museum of Natural History [Vol. LXXII

that was not clearly the direct result of some individual pathological
condition. However, one skull of Dendrohyraz. d. nigricans in the col-
lections of the American Museum is exceptional in a distinct warping of
the skull. Here the right side is longer than the left, the difference
being most obvious in the occipital region. Correlated with this asym-
metry is a difference in the upper tooth rows, that of the right side being
nearly straight, that of the left clearly bowed.

DENTAL ANOMALIES IN THE HYRAXES

Parallel to the cases so frequently encountered among the rodents,
where too the proper occlusion of the persistently growing incisors is
dependent on even wear of the tips, are hyraxes in which the upper
incisors, unopposed by wear below, grow in great ares. No specimen in
the Congo collection shows this anomaly, but the American Museum
possesses one skull of this type picked up by Dr. Chapin on Mt. Kenya
at an altitude of 14,000 feet. The specimen is that of a Procavia 7.
mackindert.

Another similar anomaly occurs in a specimen of an old male Dendro-
hyrax d. eminz in which one of the upper incisors had been lost long be-
fore death and the two lower incisors, which would normally oppose this
tooth had grown forward about twice as far as the corresponding teeth
on the opposite side. This same individual had suffered a broken zygo-
matic arch on the same side as the tooth loss, at what was probably the
same time. Most of the chewing teeth had been broken and destroyed
during life and it appears that this ancient hyrax may have died through
the accidents to which his dental equipment had been subjected.

The first permanent teeth to wear completely away are the first
molar above, the first molar below. These teeth are fully fune-
tional and well worn before the deciduous fourth premolar is shed and
before the second molar is up in position. The condition in which the
first molar is worn down to the roots is encountered in all of the genera
of hyraxes, and is not uncommon, but specimens in which the second
molar is also worn out are rare. I have encountered no skull in which
the crown of the third molar has disappeared.

SEXUAL CHARACTERS IN THE SKULLS

The sexual dimorphism of the upper incisors was established by
Thomas (1892, P. Z. S., p. 54). As he pointed out, in some species the
dimorphism is not well marked. It is, however, firmly established in

1936] Hatt, Hyraxes Collected by American Museum Congo Expedition 123

both Procavia j. lopest and Dendrohyrax d. emini where the sex of any
adult may be easily recognized by these teeth.

The size of the skull in the Congo species seems to be uncorrelated
with sex as was also recognized by Thomas for the hyraxes as a whole.

Sex differences in the skulls of the Procavia 7. lopest and Dendrohyrax
d. emini are not the same. In the latter, one observed difference is that
the skulls of the males are more massive and rugged, the ridges overlying
the roots of the incisors larger in the males, reflecting the larger size of the
teeth.

Sex differences in the skulls of the rock hyraxes of Aba (Procavia j.
lopesz) are centered about the muzzle and the incisors, reflecting a better
fighting equipment in the males. ‘Thus the heavy bony ring at the gum
line of the incisors is in the males enlarged and rough, and the muzzle is
slightly more massive. Most striking is the difference in the maxillary
root of the zygomatic arch, the outer surface of which in all adult males
is deeply concave, while in the females and young (Stage V) males it
slopes gently from the gum line to the maxilla-malar suture. For pur-
poses of sex diagnosis this is second in usefulness only to the shape of
the incisors. The mandibular symphysis further reflects the increased
strength of the male buccal region in the development of sharply de-
lineated ridges which carry forward the line of the lower mandibular
borders and pass upward to meet high on the symphysis.

In the males of this species there is the further difference that the
frontal bone is on the average flatter and broader than in the females.

Tue Aces aT WuHicH THE UPPER CANINE Is Lost

The deciduous canine is lost at different ages in different genera, as
Hahn has shown. My own figures, given below, bear out his conclusion
that in Procavia the tooth is lost before Stage V; that in Heterohyrax
it may be retained in Stage V; and in Dendrohyrax dorsalis is always
present to at least Stage IV and commonly to Stage VII.

In Procavia j. lopesi the five specimens in Stage III all retain the
upper tooth which Lataste (1886, Ann. Genova Mus., (2) IV), con-
cluded was a canine, whereas all specimens in stages older than this have
lost this tooth. The Matadi hyrax (H. chapinz) does not have this
tooth in the specimen in Stage VIII, but does in the other which is in
Stage V. The canine is retained in all specimens of D. d. emini, seven in
number, of Stages I to V and is lost in the six specimens of Stages VI
and VII. There are twenty-four specimens in Stage VIII, in twenty-

124 Bulletin American Museum of Natural History [Vol. LX XII

two of which no canine is present. In the two others, however, a canine
is retained on one side.

SOME CHARACTERS UTILIZED FOR CLASSIFICATION OF HYRACOIDEA
AND THEIR VARIATION IN THE CONGO SPECIES

CRANIAL CHARACTERS

Because Hahn’s monograph treats exhaustively of the value of most
commonly used skeletal characters it would be superfluous for me to dis-
cuss them in general, but the series which I have studied have furnished
some information not covered by Hahn or at least neglected by him.
Some of this is here recorded.

CLOSURE OF THE ORBITS

No specimen of either Procavia 7. lopest or Heterohyrax chapini in the
Congo Expedition collection has closed orbits. Every skull of Dendro-
hyrax d. emini examined has closed orbits, even in Stage I. In this re-
spect all specimens are in harmony with their generic standard. In
many other species of hyrax such constancy does not obtain.

LENGTH OF DIASTEMA

Diastema length, as I have noted above and as may be clearly seen
from an examination of the table of cranial measurements, is subject to
too great variation in equal aged individuals of the same sex, from the
same locality, to be a reliable criterion as to race. Generic differences
are well marked, it is true, and occasionally specific differences. The
length of the diastema increases with age, both actually and relatively,
in most, if not in all species.

THe CouRSE OF THE TEMPORAL RIDGES

The course of the temporal ridges in my experience, although Brauer!
believed otherwise, has proved totally inconstant in every species of
hyrax examined in series. Among equal-aged specimens of one sex of one
species (e. g., D. d. eminz) are found ridges that are parallel along the
greater part of their superior border; others strongly bowed and most
nearly approximate at the middle of the parietal; others which converge
sharply from the anterior end of the parietal to its posterior border.
For this reason I do not consider the pattern as indicative of genetic
relationship.

11934, Zeitschr. fiir Siugetierkunde, Bd. IX, pp. 198-206.

1936] Hatt, Hyraxes Collected by American Museum Congo Expedition 125

SEPARATION OF MALAR AND LACRYMAL BONES

The malar and lacrymal bones of Procavia 7. lopest usually come in
contact before Stage VIII. In only one of ten Stage VIII skulls is there
any separation, and this is but slight. Two Stage VII skulls have a gap
of about 1 mm. between the bones, while two in Stage VI already have
the bones in union. Three Stage V skulls show a gap between the two,
while two in Stage IV show it closed. In three Stage IIT skulls there is
contact of malar and lacrymal, while in a fourth these bones are sepa-
rate. Thus for the rock hyrax of the Upper Uele it seems the rule that
there is union of these bones in adults, but that the time of first contact
is exceedingly irregular and may occur very early in life.

Union of malar and lacrymal bones in the Congo tree hyrax is rare,
having been observed in only one of 24 Stage VIII skulls and in none of
the nine skulls of earlier stages. It is further evident from an examina-
tion of the table of cranial measurements of Stage VIII specimens that
the distance between these bones is on the whole fairly constant in the
species.

These observations bear out Brauer’s (1934) conclusion that the
malar-lacrymal relationship serves as a good generic character.

THE BREADTH OF THE LACRYMAL PROCESS

Brauer has used the breadth of the lacrymal process in characteriz-
ing species, but the great individuality shown in the size of this process
among series of hyraxes from a single locality suggests that this is in-
significant and useless as a criterion. In Stage VIII Procavia 7. lopesi
this process varied from 2.2 to 3.3 mm. in width, while in Dendrohyrax d.
emini the range is from 2.5 to 5.6 mm. ‘This variation embraces the
width of practically any specimen of hyrax in the Museum’s collection,
and the character is not believed to be of use in so far as the species here
represented are concerned.

POsITION OF LACRYMAL FORAMEN

The position of the laecrymal foramen was found by Brauer to be of
significance in some cases. In the Congo collection there is high vari-
ability in this structure that is uncorrelated with either age or sex, and
though this variation is far greater in Dendrohyrax d. emini than in
Procavia j. lopesi it is believed that the feature cannot at present be
considered stable in either species and hence reliable for taxonomic

126 Bulletin American Museum of Natural History [Vol. LX XII

purposes. The position of the lacrymal foramen (on one side, there
being little bilateral asymmetry) as found for these two species is as
follows:

COMPLETE COMPLETE

BELOW BEHIND

PERIPHERAL PROCESS PROCESS
Dendrohyrax d. emini 9 16 vig
Procavia j. lopest 20 if 0

A survey of all the hyrax skulls in the American Museum indicates
that all types of lacrymal foramina occur in all of the genera, but that
central foramina are more common in the tree hyraxes and marginal
foramina more frequent in Heterohyrax and Procavia.

THE VENTRAL SURFACE OF THE AUDITORY BULLAE

The ventral surface of the auditory bullae of hyraxes is fairly con-
stant in shape within any age group of a species of hyraxes, and as
different species show different forms of bullae the character has been
used (by Brauer et al.) to distinguish the species. The character, how-
ever, is limited in its usefulness for the best descriptive terms are am-
biguous and these bullae can at best be described as more or less inflated
than those of another species to which they are compared.

Dendrohyrax dorsalis latrator (Thomas)

Procavia emini latrator THomas, 1910, Ann. Mag. Nat. Hist., (8) V, p. 285.
Type locality: Batempa, Upper Sankuru River, southern central Congo.

Dendrohyrax dorsalis emint HAHN, 1934 (part), Zeitschr. fir Saugetierkunde,
Bd. IX, p. 259.

A race distinguished from D. d. emini by the white or whitish coloration of the
basal half of the body hairs.

Represented by a single, incomplete native skin, unsexed and
juvenile, obtained at Bolobo in December, 1914. The American
Museum has one other native skin of an adult, collected at Lukolela by a
more recent expedition.

These two incomplete specimens, one probably of a Stage I indivi-
dual, the other possibly of one in Stage VIII, bear out Thomas’s original
description of the race, as far as the skin areas are represented.

It would seem from the localities represented by the type and these
two specimens that the range of Dendrohyrax dorsalis latrator extends
along the southwestern border of the Congo Forest.

Dr. James P. Chapin informs me that the voice of D. d. latrator, as he

1936 | Hatt, Hyrazes Collected by American Museum Congo Expedition 127

has heard it about Lukolela, is entirely different from that of D. d. emina
as he knew it to the east. The two, he states, could never be confused by
their calls.

Dendrohyrax dorsalis emini Thomas
Plates XVIII to XXI; Text Figure 1

Dendrohyrax emini THoMas, 1887, Ann. Mag. Nat. Hist., (5) XX, p. 440. Type
locality: Tingasi, Monbuttu (2° 30’ N., 27° 50’S.), Belgian Congo.

Fig. 1. Dendrohyrax d. emini. Male, Stage VIII. Niapu, Belgian Congo.
A. M. N. H. No. 53830. Natural size.

Dendrohydrax beniensis BRAUER, 1917, Sitz. Ber. Gesell, naturf. Freunde, p. 295.
Type locality: Beni Zambo, between Kalumenda and Beni, Belgian Congo.

Dendrohyrax congoensis BRAvER, 1917, loc. cit. Type locality: Beni
(Kartoushi) [slightly north of Beni, close to the Semliki River], Belgian Congo.

Dendrohyrax rubriventer BRAUER, 1917, loc. cit. Type locality: Kalumenda,
Beni, Belgian Congo.

Dendrohyrax brevimaculatus BRAUER, 1917, loc. cit. Type locality: Lesse,
eastern Belgian Congo.

128 Bulletin American Museum of Natural History [Vol. LX XII

Dendrohyrax dorsalis nigricans Haun, 1934 (part), Zeitschr. fiir Saugetierkunde,
IBGL, IDX, {0 AZ

Dendrohyrax dorsalis enunt Haun, 1934 (part), op cit., p. 259.

Dendrohyrax d. emini is a large tree hyrax typical of the genus in cranial char-
acters and the mammary formula. It belongs to the dorsalis group of forms in
which the muzzle is largely naked in the adult animal, but differs from the typical
subspecies in the pelage which in emznz is usually more lightly colored and harsher.
In many skull characters it resembles D. d. dorsalis but differs from that race
in the fact that the interparietal-parietal sutures are closed and tbe interparie-
tal-supraoccipital suture open, the reverse of which obtains in the Gulf of Guinea
specimens.

Represented by 33 skins, 2 skeletons, 37 skulls and 5 fetuses in alco-
hol, collected as follows:

Akenge, 3 (oc), Oct. 2-18, 1913.

Avakubi, 1 (¢), Oct. 22, 1909.

Gamangui, 5 (o’"), Feb. 10-18, 1910.

Medje, 6 (52 @, 1 fetus), Jan. 21—Oct. 1, 1910.

Ngayu, 2 (o’, 1 fetus), Dec. 19, 1909.

Niangara, 3 (o’o"), Nov. 20, 1910—April 28, 1913.

Niapu, 24 (1400, 5@ Q, 2 Juv. sex?, 3 fetuses), Nov. 10, 1913-Jan. 3, 1914.

The ages are represented as follows:

STAGE MALES FEMALES SEx?
I e: 2 2
II ee Se
Ill i 1
IV hs
V 2,
VI 1
VII 4 Be
Vill 16 8
24 11 2

Specimens with skins were taken in every month except March,
June, August and September, but there is poor representation of such
specimens in all the season from March to September. This is, how-
ever, of little importance for none of the specimens were taken as much
as four degrees from the equator, and the two short dry periods in
January and July probably produce no change in the pelage of the
hyraxes.

The monthly catch of specimens was as follows: January, 2; Feb-
tuary, 3; March, 2; April) 2. July, 1; October, o;” Novemivemgar
December, 13.

1936 | Hatt, Hyraxes Collected by American Museum Congo Expedition 129

The mammary formula of Dendrohyrax d. eminz, as determined from
an examination of skins of 8 females from the Congo series, is 0-1 =2,
which formula is typical of Dendrohyraz.

THE PELAGE
Dichromatism

Dichromatism in hyraxes was first noted by Aharoni! who observed
that in the Syrian species some of the individuals were light colored,
others dark. Shortly after this, in June, 1932,? MM. Heim de Balzac
and Bégouen in the description of Heterohyrax antineae of the Hoggar
Plateau, Central Sahara, noted two well-marked color phases, but mis-
lead by an atypical combination of cranial characters they did not recog-
nize that one of the color phases of their supposedly new species was
nothing more than the earlier known Hoggar species Procavia bounhiola
Kollman, a slip which Schwarz? soon discovered.

Dichromatism, as I have found, is also strongly marked in Dendro-
hyrax d. emini, a circumstance which mislead Brauer into describing too
many forms and Hahn into recognizing too many. The common phase
of the species is the type with a broad yellow wash (Plate XX, center),
the rarer, a dark pelage (Plate XX, right). Between these are specimens
superficially intermediate, but in reality these others are worn stages
of the light and dark types. Color and color pattern in these two sorts
of animals are not precisely constant, since lighter animals have, for
example, dirty white, light yellow or orange bellies, and some less con-
spicuous features show a wide range in variation. That these color
varieties are early established is shown by two Stage I specimens of
approximately equal age, both females, one taken October 22 at Avakubi,
the other January 21 at nearby Medje. The pelages of these two, well
shown in Plate XIX, differ throughout, the bases as well as the tips of
the hair shafts being darker in the dark individual than in the light.
The diffusion of the pigment extends to belly, hands and feet.

The type specimen of D. d. emini, to judge by the color plate pub-
lished of it (1888, P. Z. S:, Plate 1), is an animal in the more common
yellow pelage. The types of D. beniensis and D. congoensis to judge by
the original description are individuals of emznz in the darker phase.

Well worn specimens of the light emznz (Plate X XI, center) are
intermediate in appearance, the nae tips to the hairs being gone, the

11930, Zeitschr. fiir Siugetierkunde, V,
21932, Bull. Mus. Hist. Nat. Paris, (2) Py? we
31933, Ann. Mag. Nat. Hist, (10) xeLIs p. 625

130 Bulletin American Museum of Natural History [Vol. LX XII

dark bases show out and present a color much like that of true ‘‘darks.”’
Worn “‘darks’’ are in turn slightly darker in general tone than their
genetic twins in fresh pelage, for in individuals of the dark type there
is a narrow subterminal band of brown which gives a grizzled appear-
ance to the coat.

One individual in the collection (Plate XX, left), No. 53819, a Stage
VIII &@ taken November 20, 1910 at Niangara, is unique in the presence
of three well-marked rusty-orange bands crossing the body transversely,
one over the shoulders, one at the level of the anterior end of the dorsal
spot (between these there is a light wash of the same color) and a third
across the back at about the level of the crests of the ilia. These bands
extend well down over the flanks but do not pass on over the belly.
Basically the individual is a normal light phase, and it might be pre-
sumed that these bars were due to some period of faulty packing or
storage of the skin, but because the orange color is exactly matched by
the hair tip color, occurring as a uniform wash over other specimens, I
am inclined to believe the unique pattern to be epidermal in origin.

Mr. Lang made some notes on the skin and eye colors of a living or
freshly-killed male brought in to him alive at Ngayu, December 19, 1909.
These notes, directly transcribed are:

Snout and skin that is visible beneath the scanty hair up to and around the
eyes, dark gray, nearly black. The tip of the lower jaw shows the same color. The
skin visible on ears, dark gray, inside pinkish. Pupil round, iris dark brown. Pads
on feet dark gray, pinkish in middle. The large elliptical place about the dorsai
gland is pinkish white.

Emin’s hyrax, diverse as are the colors and color patterns encount-
ered, does not have any difference in the pelages of the sexes, as far as
my eye can determine. Color, length and quality of the hair are seem-
ingly equal in male and female.

A careful and detailed examination of this series has failed to reveal
the slightest difference between specimens from any of the localities
represented. It is concluded that within the areas the species is homo-
geneous.

Age changes in pelage involve at least two phenomena. ‘The first of
these is the replacement of the soft woolly juvenile pelage by a coarse
coat typical of all adults of the species. A second change, progressive
through life I believe, is the falling away of the hair in the region of the
snout.

Mammals living so close to the equator as do the hyraxes in this
series are not subjected to great seasonal fluctuations in climate. In

1936 | Hatt, Hyraxes Collected by American Museum Congo Expedition 131

this area there are periods focussing in January and July in which the
rainfall is slightly lighter than in the balance of the year, but the differ-
ence is small and the ‘“‘seasons”’ short, with the result that pelage changes
are not known to occur in response to the little seasonal change. The
skins of this series were critically examined with reference to any such
possible change, but none was found.

Many specimens in the collection have a reddish-yellow tinge, par-
ticularly marked on the lighter underparts, that is due either to a soil-
ing in the original environment or to changes occasioned by grease in-
completely removed from the study skin. A vigorous application ef
benzine serves to remove much of this color, but some of it, which I
assume to be natural to the hair, remains. One striking example of a
hyrax with a coloration that suggests staining, yet proves non-remov-
able, is the yellow-banded individual (No. 53819) pictured in Plate XX.

ATTAINMENT OF FULL GROWTH

There is a range in total length of specimens with skulls in Stage
VIII from 525 mm. to 625, with the individuals scattered fairly uni-
formly between these extremes. The four with skulls in Stage VII also
lie within these limits, the lower limit coinciding with that of Stage VIII,
the higher limit lying at 535 mm. Two specimens in Stage VI are 525
and 545 mm. long, and the one individual in Stage V, is 550 mm. long.
It would thus appear that full growth in this species is attained at
about Stage V, though the greater upper range limited to Stage VIII
suggests a continuation of growth after Stage VIII is first attained.
The one specimen in Stage IV and the one in Stage III are 465 and 460
mm. long, respectively, which is distinctly smaller than any of those in
Stages V to VIII.

BEHAVIOR

Field notes made by Mr. Lang at Ngayu on December 19, 1909,
Medje, January 22, 1910, and Gamangui, February 18, 1910, shed some
light on the habits of the species. These notes combined for brevity and
slightly altered in wording where necessary, are as follows:

GENERAL BEHAVIOR.—A male brought in alive by the pygmies, moved very
slowly on the ground when undisturbed, but would rush rapidly for a yard or two
towards a stick thrust at it, or a person that would approach. At such times it would
turn its eyes so far backwards that only the yellowish white membranes in front or
in back could be seen, and would also erect the hair along its back and that on its
head. This little beast understands how to command the respect of all.

132 Bulletin American Museum of Natural History [Vol. LXXII

Voicr.—The animal, when disturbed, as described above, would make a short,
abrupt, pig-like grunting noise. It continually ground its teeth (seemingly the
molariform series) producing a loud noise by these movements. When taken by the
neck it would make a soft whistling noise, rapidly repeated.

The tree hyraxes in the forest cry or rather howl for ten minutes or even half an
hour with practically no interruption, repeating one long-drawn howl! after another.
Sometimes, however, one howls as rapidly and as strongly as though it would defy
all competition. One would rather attribute the call to a cat-like animal on account
of its peculiar sound. The animals start about 9:00 p. mM. and cry particularly about
10:00 F. m., though they may be heard even as late as 2:00 A. M.

These night calls of the tree hyrax may be heard a half an hour’s journey
through the forest. No more than a single animal was heard calling in the same
place. Apparently only males howl, as for a long time only males were brought in,
and in the total collection the males are more abundant.

The calls were heard in many places, the first time in Batama, September 15,
1909.

Foop AND FrEEpING.—The pygmies and other natives at Ngayu claim that the
tree hyrax descends the trees at night and feeds on the ground. The stomachs of
three byraxes taken at Gamangui contained chewed up leaves, all of the same
species of trees. One of these stomachs also contained three bees in a ball of hair,
all practically intact.

Native Caprure.—The tree hyrax, although common all over the forest, is
difficult to procure, as it does not start howling until after night-fall, and then the
natives are afraid to wander about in the forest on account of leopards.

These hyraxes keep to the same place with great persistence, as they are nearly
always heard in the same direction. The natives even claim that one keeps to the
same tree for a certain time, which seems probable, as one was heard howling every
night for about six days in at least the same clump of trees.

The natives secure the hyraxes by locating the tree upon which the animal
is howling. In the morning they climb the tree and search for it, cut down the tree
in order to secure the animal, or watch the animal closely until they find some op-
portunity to capture it. One specimen was taken from a dead hollow tree.

Heterohyrax chapini (Hatt)
Plate XXII; Text Figure 2

Procavia chapini Harr, 1933, Amer. Mus. Novitates, No. 594, p. 1. Type
locality: ‘‘Loadi Hill, 5 kim. SW. of Matadi, Bas Congo.”

Heterohyrax syriacus bocagei HaHN, 1934 (part), Zeitschr. fiir Saéugetierkunde,
Bde PX p3283.

A pale drab, coarse-haired bush hyrax with a well-marked light yellow dorsal
spot. The mammary formula is 0-2=4. The cranial characters are those of the
heterohyraxes except that the skull is larger, the muzzle longer, the dorsal profile
flatter and the basisphenoid more elevated than in H. syriacus and its subspecies.

Represented by two females, one adult, one juvenile and two em-
bryos from the adult. These were collected by James P. Chapin near

3Dr. Chapin (field notes) describes their cries as ‘‘agonized reiterated screeching.”’

Collector’s Measurements of Dendrohyrax d. emini

SKULL
Locanity SEx NuMBER TotTau Tatu Foor EAR STAGE
Akenge of * 53803 605 25 79 ay VIII
= oF 53804 545 35 85 31 VI
oi of 53805 550 30 82 35 V
Gamangui 53808 600 20 84 WADULE
2 ot 53809 620 22 82 WADOL
5 of 53810 TBY5) 21 80 VIII
i‘ of 53811 580 a 82 VIII
Ngayu CF 53818 550 20 80 te VIII
Niangara of 53819 525 20 78 30 VII
os of 53821 520 22 80 29 Bale
Niapu ii 53822 538 18 MD 35 WATE
se of 53823 540 20 87 32 WUE
a oF 53825 555 20 86 31 VII
a ow 53829 525 15 ade 35 VIII
ie oi 53830 562 18 a 32 VIII
3 of 53831 605 25 88 36 VIII
ee oH 53832 545 28 84 31 VIII
yy oi 53833 535 , DES 80 32 Wael
= of 53834 525 25 85 31 VI
of of 53835 525 25 81 31 WADUE
of 53836 595 25 88 34 WADI
a of 53837 575 25S 83 35 VIII
¥ of 53842 550 20 WE 29 VII
i of 53844 595 20 82 33 WADUL
Average CH ot 558.3 20.8 78.5 S2P VII-VIII
Minimum oo 520 15 75 29
Maximum oo 620 30 88 36
Medyje Q 53814 575 18 82 de VIII
i Q 53815 585 25 85 32 \WA00L
~ Q 53817 635 25 80 33 WAQUE
Niapu Q 53824 580 20 82 32 VIII
" Q 53826 565 20 85 30 WA00L
* 2 53828 590 18 82 32 VIII
. g 53838 585 25) 85 35 VIII
o 2 53840 585 15 85 34 VIII
Average 22 585 20.7 83.2 32. VIII
Minimum QQ 565 1S 80 30
Maximum @Q @ 635 25 85 36
Average Cl ct 2-2 565 ZS 79.7 32. VII-VIlI
Minimum o'o' @ 9 520 15 75 29
Maximum o'o' 2 Q 635 30 88 36

134 Bulletin American Museum of Natural History [Vol. LX XII

Matadi, December 27, 1914. The adult and juvenile are preserved as
skins with skeletons, the embryos are preserved in formalin.

A novelty of the collection was Chapin’s bush hyrax, secured as near
to the mouth of the Congo as any hyrax is apt to occur. The reduced
number of mammae (if the type specimen is indicative) set this species
apart from others of the genus, and its nearest neighbor, Heterohyrax
bocagez, is quite different in its longer, thicker pelage, smaller size,
broader teeth, proportionately broader skull, elevated supraorbital
ridges, shorter muzzle and flatter basicranium. (See Plate X XIT.)

Fig. 2. Heterohyrax chapini. Female, Stage VIII. Type. Natural size.

Collector’s Measurements of Heterohyrax chapini

SKULL
LOCALITY SEX NUMBER ToTtaL Tat Foor EAR STAGE
Matadi Q 53800 a 12 68 35 VIII
e Q 53801 430 10 60 30 Vv
Average 2 © | 476 1] 64 32.5

1936] Hatt, Hyraxes Collected by American Museum Congo Expedition 135

Procavia johnstoni lopesi Thomas
Plate XVIII; Text Figure 3

Procavia lopest THOMAS, 1907, Ann. Mag. Nat. Hist., (7) XIX, p. 520. Type
locality: Kodja Hill, Gaima Range, Monbuttu, Congo.

Procavia ituriensis BRAUER, 1917, Sitz. Ber. Gesell. naturf. Freunde, p. 303.
Type locality: Ituri, Belgian Congo.

Procavia johnstoni lopesi Haun, 1934, Zeitschr. fiir Saugetierkunde, Bd. IX, p.
293.

A large-toothed, coarse-haired rock hyrax apparently closely related to P. 7.
matschier. The general color is Raw Umber to buffy brown. The crown is black
and this shade occurs down to a sharply drawn line below the eye. The dorsal spot
is broad and a deep Colonial Buff in color.

Represented by 23 skins, 6 skeletons, 17 skulls and 2 fetuses in aleo-
hol, collected as follows:

Aba, 28 (8 oo, 18 9 2, 2 fetuses), July 16, 1911—January 2, 1912.

Faradje, 1 (co), April 1, 1911.

Vankerckhovenville, 1 (co), August 7, 1910.

The ages represented may be summarized in the following table:

STAGE Mates FEMALES
Zalt

II te . bes
III 3 2
IV : 1 or
V 2 il
VI 2
VII at 2
VIII 4 6

It is thus apparent that the series of fully adult specimens is not
large.

The seasonal distribution of specimens is only fairly satisfactory, the
animals with skins being secured as follows: January, 3; April, 1;
July, 3; August, 1; December, 15.

DISTRIBUTION

Procavia 7. lopesi was first made known by the description of Thomas
who had two specimens collected by Boyd-Alexander on Kodja Hill, de-
scribed by this noted explorer as “‘ directly in back of the Gaima Range,”’
which in turn lies on the left bank of the Kibali, below Vankerckhoven-
ville. The Congo Expedition specimen labeled Vankerckhovenville
was shot by Dr. Chapin on a hill behind Mt. Gaima which is, in all
probability, the Kodja Hill of Boyd-Alexander, though the collector is

136 Bulletin American Museum of Natural History [Vol. LXXIT

not perfectly certain on this point. The other specimens in the Congo
collection are from hills in the vicinity of Faradje and Aba.

THe PELAGE

Procavia j. lopest appears to be one of the least variable of hyraxes as
regards its pelage. From early youth (Stage III) to old age the color
pattern, the color and the character of the pelage remains almost un-
changed.

Fig. 3. Procavia j. lopesi. Male, Stage VIII. Faradje, Belgian Congo.
A. M. N. H. No. 58799. Natural size.

There is a very slight diminution of the orange tint of the throat and
belly between Stages III and VIII, and specimens of older individuals
show a somewhat greater restriction of black pigment in the region
of the ears, but beyond this, age leaves little mark upon them.

From hair characters I am unable to distinguish the sexes. If there is

1936 | Hatt, Hyraxes Collected by American Museum Congo Expedition 137

an annual cycle of pelage change it is not indicated in this series which
contains specimens scattered fairly well over the year. ‘The series can-
not be expected to show geographical variation as the three localities
represented are close together—about one hundred miles apart in the ex-
tremes.

ATTAINMENT OF FULL GROWTH

The smallest specimen in Stage VIII measures 480 mm., the longest
585. Of the two specimens in Stage VII, the smaller measures only
three mm. less than the smallest in Stage VIII, the larger well within the
limits of variation of the above. ‘The single skull in Stage VI is from a
specimen longer than either of those in Stage VII, while three in Stage
V are from specimens practically within the limits set by the two in
Stage VII. The one skull in Stage IV is from a specimen 15 mm.
longer than the smallest animal with a Stage VII skull. Two skulls in
Stage III are from animals 378 and 382 mm. long. Thus, animals fully
adult as measured by the attainment of function of the complete denti-
tion range in size between limits that are within five mm. of embracing
the range represented by all specimens down to the stage in which the
second upper molar is only partly up. Animals in Stage IIT (M! fune-
tional, M? below the level of the bone) are distinctly smaller. It would
appear from this that full growth in this species is attained at about the
time of the eruption of the second upper molar, which contrasts with the
situation suggested by the Congo Expedition collection of tree hyraxes
(Dendrohyrax d. emini) where full growth seems not to be attained
before the period of full function of the second upper molar (Stage V).
The period of time between the eruption and the functioning of M? may
not be great, however, and actual differences in growth pattern between
these two species may be very slight, or even nonexistent, for the series
with which I have worked is too small to furnish statistical proof.

BEHAVIOR
Notes made by Mr. Lang at Aba, December 18, 1911 are:

The colonies on these rocky hills are rather small, three or four being the usual
number of animals. They are, however, found on every rock, the cracks and boulders
of which offer sufficient shelter to them. The largest number found in one colony was
fourteen, but in this case the rock is exceptionally large and the hyraxes from the
whole rock assemble at one side to bask in the sun. On this rock is a place where the
water has collected. The hyraxes evidently wade into the water (about 5 inches
deep) to chew off the sprouts of a flowering water plant.

138 Bulletin American Museum of Natural History [Vol. LX XII

The hyraxes leave their burrows or ledges towards three in the afternoon and
probably feed throughout the night. At eight in the morning and slightly later
they bask on their ledges, but are always watchful. After ten o’clock they are not
seen out.

These animals are very shy and make a low barking (somewhat guttural) sound
at the approach of danger. They run swiftly along and upward on the strongly
inclined rocks where it would be impossible for any man to get a foothold.

These hyraxes evidently give birth to their young the end of December or the
beginning of January.!

The hyraxes nibble off the fine grass growing between the rocks. This grass
when chewed or rubbed has an aromatic smell. Their stomachs contain, chiefly, this
grass finely chewed up. There are many other plants fed upon, also the ordinary grass,
especially that sprouting freshly after being burned out in the beginning of the dry
season. ‘The natives assured me that the hyraxes are often found far from their
rocks, feeding on young sprouting grass.

Collector’s Measurements of Procavia j. lopesi

SKULL

LOCALITY SEX NuMBER Totat Tait Foot EAR STAGE
Aba of 53779 515 20 ial 33 VIII

ee of 52780 490 Pall a 29 V

cy of 53784 556 20 75 Di VIII

ze of 53786 475 20 65 Dil Vv

ne of 53791 580 24 ie 35 VIII

om of 53797 545 22 70 35 VIII
Faradje of 53799 575 11 76 33 VIII
Average Ci of 534 19.7 Walia 31.3 VII-VIII
Minimum ot! CH 475 11 65 2
Maximum ol ot 580 24 76 30
Aba Q 53776 570 ee al 38 VIII

a 2 SA MU 566 20 val 36 VIII

s 2 53781 520 22, 67 32 VII

a Q 53783 526 21 il 31 VI

ns Q 53787 545 2, 68 32 VIII

2 53788 525 Pal 67 32 V

oH Q 53793 585 24 76 32 VIII

of Q 53795 A477 20 66 33 VII

Ms Q 53796 480 18 ail 37 VIII
Average © © yA, ileal 69.7 33.6 VII-VIII
Minimum Q A4T7 18 66 31
Maximum Q 585 24 76 38
Average oo 22 533 2045 70.5 32.6 VII-VIII
Minimum oS g 475 11 65 27
Maximum | Q 585 24 76 38

1 A conclusion scarcely justified by observations made over only 7 months of the year, and on
few adult females.

1936 | Hatt, Hyraxes Collected by American Museum Congo Expedition 139

The natives (Logo) try to catch the animals with a noose, or a number hide
late in the afternoon at the rocks where the hyraxes have been observed before in
sufficient numbers. After the hyraxes are out the natives suddenly start howling
and as the animals run back to take refuge in their holes many of them are stoned
This is possible only near places where young short grass is abundant. ‘The natives
are very fond of eating these hyraxes.

Cranial Measurements of Congo

x a0)
3 =| = x
be ae = z : ; : 3 : :
a & oe = Bait eer rere Be) El q 2
do Z Boe Boe Be ee ere
‘S) _ & iS rm z a 3 4 a eI ‘3 - mo =
ae SS a Oe 9S m8. (2 be ae eee
5H 1A BS fa o4 5m S aoe se Be 2 oy
les < oD © O N MD 4 O an 4
Dendrohyrax dorsalis emina
Akenge 53803 oi 112.5 112.3" 63.3 3070° 32.5 235) ealGaoeeeZORG
Gamangul 53808 of 112.0) 112"0"" 6326" 31.9". 3120) 2a 20a eee On)
es 53809 of 11958 109.8" 6723) 322.0) {2743S 2oRomeeOn ?
es 53810 of UISey WNOses @HeO) Sus ? i sere) «Oa
a 53811 ef 111.8 109.3 66.1 935.0 30-08 2350ne LGR ORO
Ngayu 53818 of 119.7 114.2. 68.0 31.8 27.7 S232 SsOmeeeOre
Niapu 53822 ot 10655" 106.5 ~~ 62.3 31.0 29.0) 220 "aR Ge anos
i 53823 of {14.5 114.0) 63.2' 29.2 3005 “230 Sareea len
at 53829 Ci! 11458 11405" ~ 63.4- 32.02 (27-0 isan ?
2 53830 of 110.4. 109.5 58.8 32.4 24.8 20s00 Toe aeeezoet
ss 53831 a 115:0- 1138.8 64.6" 32°58 311 Qos alee
sf 53832 ow 107.3 105.7 61.3 28.7 - 2575 Zio ease Onemcerr
re 53835 ot 110.0. 109.1 ~°62:.8° 29.0 2556, 924 lee
oy 53836 of WS WIG. OS “Bl ? a iid aecons
Sy 53837 ow 110.0 110.0 --65.2 :380.2 32.1 (25568 Om Omemmion >
cs 53844 of 1138-2 -111.2 61.7 3025 S12 aly eH)
Average o'co 112.7 111.4 64.0 31.2 28.8 22.9 Lio Z0r3
Medje 52120 2 109.5 109.5 54.5 (31.5) “24567 Oh alos ao
is 53814 2 106.0 105.6 “5921 3067 27-7 2203 ono n
a 53815 fe) 108.5 “10822 5626 30. 7 > S025) 19 oes
Niapu 53824 Q 105.3 104.5 53.8" 31.0 2556) Geo lice O ees aoe
a 53826 2 104.9 104.9 °58.6 30.0 24.67 2005s ibe ORO
gt 53828 2 10955 109-4 58.1% 32-3 25.07 20 esas
S 53838 2 107.2 106.6 58.1 °28°8 29.7 SOR alse Ome teel
a 53840 2 114.0 118.7 59.8 (29.2 30.8 22253 Ge Omeeeilen
Average 99 108.1 107.8 57.3 30.5 27.3 2080) S353 Or4
Average o'? 111.2 110.2. 61.8 30.9: 28.3 (2158) 16-35 20-0
Heterohyrax chapini
Matadi 53800 2 95:0 94:6 50.3 31:8 22.0> 2IMOn ako meeleed
Procavia 7. lopest
Aba 53779 i 97.5: —96.5 55.4 3158 27.5) QING Ree oe elon
a 53791 of 96.8 - 95.9 (57.3 33:2 (24.3% Di Reine Oem
a 53797 of 96.7 938.7 56.7 34°76 (25.022 aloes
Faradje 53799 oF 98.7. 98.5 62.3. 34:0 26:.'7 > 2455 ete:
Average oo 97:4 96.1 57.9. 33.4 25,9) 522eIh Selo HOmetie9
Aba 53776 2 103.0. 10225: 57.2, 32:0) 2720) 2205Seel eae)
oi 538777 2 98.8 9557 57:8 34.2 24.4) 520 Sie sill Semele,
af 53784 2 96.0 > 96.0 56.4 32.4 24.2°- 2100) Se omeellowd
ns 53787 2 94.5 93.5 56.3 31.2 25.6) 2220 ie eelon0
sf 53793 Q 96.0:2°°94.5 9 5720 382.8 24.3." 21 a2 aes
or 53796 2 92.0. (89.8 51.8 30.8 24.5 1Os2 OsGilGe2
Average 92 9 96.7: 95.3. 56.1 .-32..2 »2550. Zin Omueizer 16.8
Average o@Q 97.0" 95.7... 56.8 (32.7 25.3 21NSo) 24a

Expedition Procaviidae (Stage VIII)

I-Wd HIONaT]

e-JN LHDIGH

I-WW Hii AA

G-INNd HLGIM

HLONG'I UVTOPAL
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aITd1dNVN
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AVSSOd 'IVHOdWAT,

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“IXOUddV LSASOT/)

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GONVISIC LSVO'T

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WUOCAS
IVINOWdA HALO NAT

VINALSVICT

3.3 4.5
4.2 4.3

ORAS NO SZ ez OP DOSE oe AOR ou eee Gol
eves (f

5.8
2.3

29.5
4.3

36.4

15.3

0) AN) > Casa =a) kell eee

6.4

18.4

sand | 29.3

19.0

6.5

10,9 a 4s.8

29.0 Ib 8) 20.7

32.5

21.3

Pe Pope Lunt
2.6 4.6
3.3 4.5

15.3 ORS So OZ Ane 9G. 14 OO
11.8 5.2
3.4 3.8

Day)

3.8
3.7

29.8
3.3

34.9

MERLE

6.5

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30.5 23.9 IN).

38.8

18.4

Ve Ins Oil Zabe0) = 1d.0)
Ong

14.7

30-0 28.1

15.6

12.5

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LOT,

19.5

18.5

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3.2

27.9

2.5 4.3

SSO OZ le Or on eae ell
aa) ).33

21.5

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16.8

BOO P5724 Reyes 4b) WG) ALO W)ieeh athe) 9 4bo (574

16.1

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Mel aeo8

3420) 227.2 4.6 14.4 Dads PANO) ete)’ sy areietss Abs es)
(Hts “ber

16.4

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13.2 5.0

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5.0

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4.0

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16.8

6.9

42.3
39.5

21.6 104.0
20.3

18.6

34.2 30.0

18.6

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10.0

16.3

a2 59 7 29).0

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sone 2o. 0 4.0 gle NO. PAL | Oiigen- aw il ab re) al)
6 ALO G8

18.4

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17.6

3.6 4.0
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2.7 4.3
3.7 4.2
4.2 5.3
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Pholly. Beet)
3.0 4.6

Sono 28.1 ~4./ sss Os WOSO > SILO Sih) Ares) = WO)

ees)

Seay 2962 28 N45 NOS Ao Cage raleisiy —-4b56) * Wea

15.8

6.0
6.1

ZonOme ss 10.5 S500 2051 SOn oO ote On ene:
4.6 4.1

33.2

18.4

Yeh te - SEA BD
40.0

10.0

15.6

32.4 27.6

iWe9

6.2
6.6
6.4

6.2

4.6
4.6
4.6
5.0

Sik A
4.5

20.0

sori 292. 4.2 14.5

15.2

BORO 2859 ~ 8.6 477 NO) = Asay E7450) =) est)

16.0

a O eel Ri O25 00 O89

10.5

17.3

36.8

16.5

Soll 4020

21.3

B02 =) 2.4, 4.5 20.3

19.6

6.2

34.9 28.3 3.9 W556) 9206S OR. OE Os, 7392

sont) ° 28.8 3.8 16.1 OF OF ZOE OF 9448459 Se 4 On 10S 2a dao

17.4

SOO 2On i TART 222, So UD.) CHE AW) DAD = Dob ow) Bok!

16.2

Auiote: | Synch tele (OG 7 Bac)
8.0

85.0

16.4

37.6

13.5

5.3 3.0

UG thd) G40) abe)

led

o1.2

12.5
12.0

36.5

Bone, 12478 Ono 3.7 Boll Mere feQaay 4h) EN) a ee Bell
Uf k3

13.4

O72, Sha2y 40.98 456 Doth Wall
4.5 3.6
4.1 4.2

74.72

S010) 2e5- | On1Z 7.6

12.8

8.5
8.2

U5 CO, eee) 1 5)5(0)
Opell

6.0
2.5

39.5

15.0

85.2 41.8

16.0

5.5

3.5
i) Bia)

Doe) Ded

Aods ME) cea DEO

4.0

4.2

10.5

8.3

4.8
5.0

42.7
4.5

82.0

16.1

orl

Foe
8.0
8.1

40.7

86.2

eS
1.5
33

5.7

39.0

12.8

Ae le a0)

ef) Ose

80.5 39.3

16.0

3.0
4.3

NOR L

84.7 40.9 4.8

37.1

11.6

4.6 3.4

8.0

85-8 40.9 4.7

16.0

37.0 24.7 0.04 6.0 2.9

1221

Puatrt XVIII

Fig. 1. Dendrohyrax d. emint.

Fig. 2. Procavia 7. lopesi. Female, Aba, Belgian Congo. Contrast this species
with that above for distribution of hair on the face, and apparent length of hair on the
back.

Specimens in the flesh. Herbert Lang, photographer.

BuLuETIN A. M. N. H. Vou. LX XII, Puate XVIII

PratE XIX
Light and dark phases in juvenile Dendrohyrax d. emini. The light (A. M. N. H.
No. 53806) is a female taken at Avakubi, October 22. The dark (A. M. N. H. No.
53812), also a female, was taken at Medje, January 21.

Vou. LXXII, Puate XIX

Buuuetin A. M. N. H.

PLATE XX
Color phases in adult Dendrohyrax d. emint. The center specimen (A. M. N. H.
No. 53836, ad. o’, Niapu, December 17) is an example of the common blond phase.
To its right is a specimen (A. M. N. H. No. 53821, ad. o, Niangara, April 28) in the
dark phase, and on the left another (A. M. N. H. No. 53819, ad. o, Niangara, Nov.
20) basically blond, but exhibiting darker transverse bands of orange color.

Buuuetin A. M. N. H. Vout. LXXII, Puatre XX

Puate XXI

Color patterns due to wear in the pelage of Dendrohyrax d. emint. The left
(A. M. N. H. No. 53836, ad. o, Niapu, December 17) is a light phase with unworn
pelage. In the center is a specimen (A. M. N. H. No. 53815, ad. 9, Medje July
27) with greatly worn pelage, in which over a broad area the dark bases of the hair
show through. The specimen on the right (A. M. N. H. No. 53828, ad. 2, Niapu,
December 1) is an example of the dark phase in which the pelage is worn about
equally to that in the center.

Vou. LXXII Puate XXI

N. H.

BuuueTIn A. M.

JPiNaiD) SOUL
Skulls of Heterohyraz s. bocaget (above: A. M. N. H. No. 80601, ad. 9, Lubango,
Angola) and Heterohyrax chapini (below: type). Contrast the length of muzzle, the
height of the supraorbital ridge, the basal eminence.

BULLETIN

AG MIE

N. H.

Vou. LXXII, PLuatrs

XXII

ag ep

oY ae ee ay

$
‘y

a’
IF

iin

3 9088 006/2 1773

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